Entomological Review of Japan

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Vol. 60, No. 1 - April 30, 2005

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Em. Rev. Japan , 60 (1): 1-7, April 30, 2005

Records of Donaciinae from Primorsky Province in 2004,
with Notes on the Distribution of Plateumaris shirahatai Kimoto
(Coleoptera: Chrysomelidae)

Masakazu Hayashi

Hoshizaki Green Foundation, 1659-5, Okinoshima, Sono, Izumo, 691-0076 Japan
E-mail: hgf-haya@green-f.or.jp

and

Osamu Tominaga

A-312, Shibatsuji-cho 4-1-15, Nara, 630-8144 Japan

Abstract Eight species of donaciine beetles taken in 2004 from south Primorsky, Far Eastern
Russia, are recorded, with the first record of Plateumaris shirahatai Kimoto from the conti¬
nental region.

Hayashi (2001, 2002) reported collecting records of 11 species of Donaciinae from southern
Primorsky Province, Far Eastern Russia including the first records of Donacia bicoloricornis
and Donacia vulgaris from the province.

We visited the region again in May, 2004 and collected eight species in two genera:
Plateumaris shirahatai , P. roscida , P. weisei , Donacia aquatica , D. flemora , D. ochloreuca , D.
vulgaris and Donacia sp. Among them, P. shirahatai previously known from Sakhalin and
Japan (Hokkaido, Honshu) is commonly found from marsh in the region.

We gratefully acknowledge to Dr. Victor N. Kuznetsov (Institute of Biology and Soil
Science, Far Eastern Branch of Russian Academy of Science, Vladivostok) and Mr. Shigehiko
Shiyake (Osaka Museum of Natural History) for supporting our field survey.

Plateumaris shirahatai Kimoto

(Figs. 1,2, 11)

Records. 4c? <?, l£, pond, 4 km E of Romanovka V., Bolshoy Kamen Town, Primorsky, 15. V.
2004, M. Hayashi leg.; 5<?<?, 1 £, pond, 4 km W of Novonezhino V., Bolshoy Kamen Town, Primorsky,
15. V. 2004, M. Hayashi leg.; 3c?c?, ditto, O. Tominaga leg.; 5c?c?, pond, 2 km WSW of Gorno-
tayozhnoye V., Ussuriysky District, Primorsky, 16. V. 2004, O. Tominaga leg.; 5 c? £?, 4-? pond, 3 km
NW of Kaimanovka V., Ussuriysky District, Primorsky, 16. V. 2004, M. Hayashi leg.; 13«?c?, 4-¥-£,
ditto, O. Tominaga leg.; 1 £, pond, 5 km WNW of Kaimanovka V., Ussuriysky District, Primorsky, 16. V.
2004, O. Tominaga leg.; 7c?c?, 1 marsh, Zarechnoye V., Ussuriysky District, Primorsky, 17. V. 2004,
M. Hayashi leg.; 18 c? c?, 8? ?, ditto, O. Tominaga leg.

2

Masakazu Hayashi and Osamu Tominaga

Figs. 1-2. Plateumaris shirahatai from south Primorsky-1, Habitus, male; 2. endophallus.

Allow indicates notched median process. Scale bar = 0.5 mm.

Host plants. Adults were collected on the flowers of Carex spp.

Remarks. Plateumaris shirahatai was described from Yamagata Prefecture, Honshu,
Japan (Kimoto and Hiura, 1971). It resembles a transpalaearctic species, P. sericea Linnaeus.
It had been known as a Japanese endemic species (Kimoto, 1983), but recorded from south
Sakhalin (Mikhailov and Hayashi, 2000), and also commonly found from several localities in
south Primorsky. It is quite within the bounds of possibility that P. shirahatai is a synonym of
other Palaearctic species (for example, P. obsoleta Jacobson) because its identification is very
difficult based on external morphology.

Distribution. Primorsky, Sakhalin; Hokkaido, Honshu.

Plateumaris roscida Weise

(Figs. 3-5)

Records. 1 <?, 1£, marsh, 1km E of Mt. Sestra, Partizansky District, Primorsky, 14. V. 2004, M.
Hayashi leg.; 4<?c?, ditto, O. Tominaga leg.; 3<?c?, pond, 2 km WSW of Gornotayozhnoye V.,
Ussuriysky District, Primorsky, 16. V. 2004, M. Hayashi leg.; 18c? <?, 1?, ditto, O. Tominaga leg.; 6<?
d\ pond, 3 km NW of Kaimanovka V., Ussuriysky District, Primorsky, 16. V. 2004, M. Hayashi leg.; 6 c?
d\ I-?-, ditto, O. Tominaga leg.; lc?, pond, 5 km WNW of Kaimanovka V., Ussuriysky District,

Donaciinae from Primorsky Province in 2004

3

Figs. 3-10. Donaciinae from south Primorsky.-3-5, Plateumaris roscidci (3^1, male; 5, female); 6-7, Donacici

ochloreuca (6, male; 7, female); 8-9, D.flemora (8, male; 9, female); 10, D. vulgaris , female.

4

Masakazu Hayashi and Osamu Tominaga

Primorsky, 16. V. 2004, M. Hayashi leg.; 1 <?, ditto, O. Tominaga leg.; 12<? <T, 2? marsh, Zarechnoye
V., Ussuriysky District, Primorsky, 17. V. 2004, M. Hayashi leg.; 14 c? <?, 5-?- ditto, M. Hayashi leg.
Host plants. Adults were collected on the flowers of Carex spp.

Remarks. In some specimens, the femora are entirely rufous and partly dark-colored (Fig. 4).

Plateumaris weisei Duvivier

Records. 1?, marsh, 1km E of Mt. Sestra, Partizansky District, Primorsky, 14. V. 2004, M.
Hayashi leg.; 1 <?, ditto, O. Tominaga leg.; 1 <?, pond, 3 km NW of Kaimanovka V., Ussuriysky District,
Primorsky, 16. V. 2004, M. Hayashi leg.; 1 marsh, Zarechnoye V., Ussuriysky District, Primorsky, 17.
V. 2004, O. Tominaga leg.

Host plants. Carex spp.

Donacia ( Donaciomima ) aquatica Linnaeus
(F ig. 12)

Records. 1 d\ 1 ?, pond, 4km E of Romanovka V., Botshoy Kamen Town, Primorsky, 15. V. 2004,
M. Hayashi leg.; 1 <?, ditto, O. Tominaga leg.; 8c?<?, 3? £, pond, 4 km W of Novonezhino V., Bolshoy
Kamen Town, Primorsky, 15. V. 2004, M. HayashI leg.; 3 <? <?, 4? ditto, O. Tominaga leg.; 1 <?, pond,
2 km WSW of Gomotayozhnoye V., Ussuriysky District, Primorsky, 16. V. 2004, M. Hayashi leg.; 1 £,
ditto, O. Tominaga leg.; 4<? <?, 7-?- pond, 3 km NW of Kaimanovka V., Ussuriysky District, Primorsky,
16. V. 2004, M. Hayashi leg.; 25 c ?c?', 18, ditto, O. Tominaga leg.; 3c?c?, 1 pond, 5 km WNW of
Kaimanovka V., Ussuriysky District. Primorsky, 16. V. 2004, M. Hayashi leg.; 4c?c?, 8■?-¥-, ditto, O.
Tominaga leg.; 3(?<?\ 4££, marsh, Zarechnoye V., Ussuriysky District, Primorsky, 17. V. 2004, M.
Hayashi leg.; 9<?d\ 7-?- £, ditto, M. Hayashi leg.

Host plants. Adults were collected on the flowers of Carex spp.

Donacia ( Donaciomima) flemora Goecke
(F igs. 8, 9)

Records. 1 pond, 4 km E of Romanovka V., Bolshoy Kamen Town, Primorsky, 15. V. 2004, O.
Tominaga leg.; 1 c?, 1?, pond, 4 km W of Novonezhino V., Bolshoy Kamen Town, Primorsky, 15. V.
2004, O. Tominaga leg.; 1 c?, 1 pond, 3 km NW of Kaimanovka V., Ussuriysky District, Primorsky, 16.
V. 2004, M. Hayashi leg.; 1 c?, 2-?- ditto, O. Tominaga leg.; 1 <?, 4-¥- marsh, Zarechnoye V.,
Ussuriysky District, Primorsky, 17. V. 2004, M. Hayashi leg.

Host plants. Carex spp.

Donacia (Donaciomima) ochroleuca Weise
(F igs. 6, 7, 13)

Records. 9<?c?, 8-¥- ?, marsh, 1 km E of Mt. Sestra, Partizansky District, Primorsky, 14. V. 2004,
M. Hayashi leg.; 11 £ S', 7-?- ditto, O. Tominaga leg.; 2cT <?, 1 •¥■, pond, 3 km NW of Kaimanovka V.,
Ussuriysky District, Primorsky, 16. V. 2004, O. Tominaga leg.; 1 <?, pond, 2 km WSW of Gorno-
tayozhnoye V., Ussuriysky District, Primorsky, 16. V. 2004, O. Tominaga leg.

Host plants. Carex spp.

Donaciinae from Primorsky Province in 2004

5

Figs. 11-14. Donaciinae from south Primorsky.- 11, Plateumaris shirahatai on flower of Corex sp.;

12, Donacia aquatica on flower of Corex sp.; 13, D. ochloreuca on leaf of Corex sp.; 14, D. vul¬
garis on leaf of Carex sp.

Remarks. Askevold (1990) recognized that Donacia ochroleuca was a synonym of D.
fennica Paykull, but Medvedev (1992) treated D. ocreleuca as an independent species.

Donacia {Donaciomima) vulgaris Zschach
(Figs. 10, 14)

Records. 1 <?, pond, 4 km W of Novonezhino V., Bolshoy Kamen Town, Primorsky, 15. V. 2004, S.
Shiyake leg.; 1 marsh, Zarechnoye V., Ussuriysky District, Primorsky, 17. V. 2004, M. Hayashi leg.

Donacia {Donaciomima) sp.

(Figs. 15-19)

Records. 1 <?, 2£ pond, 3 km NW of Kaimanovka V., Ussuriysky District, Primorsky, 16. V.
2004, O. Tominaga leg.

Remarks. This species is characterized by robust body and antennae, metallic legs, and
glabrous pronotum (Figs. 15 and 16). Median process of endophallus is short and gently arched

6

Masakazu Hayashi and Osamu Tominaga

Donaciinae from Primorsky Province in 2004

7

towards underside (Figs. 18 and 19). We can not identify it with any species known from the
Far Eastern Russia at present.

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# • Stk \%: vj (2004^) xfy?

4 - 2004^7 ft izu

X 4 £ fc^-c§£.

yy ^ 9 ^ X 9 ■' "'■U'y Plateumaris shirahatai Kimoto ^ P. roscida Weise, Donacia

aquatica Linnaeus ItZYMcOtZ-tX'#- < b- bfltz. vyJ'9 5 ->(±^5^, -tjv\U >

ifc. %bKtz7 $*>%&#}&&&b, ^mx[zm

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References

Askevold, I. S. 1990. Reconstructed phylogeny and reclassification of the genera of Donaciinae
(Coleoptera: Chrysomelidae). Quaestiones Entomologicae , 26: 601-664.

Hayashi, M., 2001. Donacia bicoloricornis Chen from Far East Russia, with records on several Donacia
from Primorskiy Province (Coleoptera: Chrysomelidae: Donaciinae). The Entomological Review of
Japan , 56: 63-66.

-, 2002. Records on Donaciinae from Primorsky Province in 2002, with taxonomic study on

Donacia knipotschi Jacobson (Coleoptera: Chrysomelidae). The Entomological Review of Japan, 57:
197-202.

Kimoto, S. and Hiura, I. 1971. A List of the Chrysomelid specimens preserved in the Osaka Museum of
Natural History , III (Insecta: Coleoptera). The Bulletin of the Osaka Museum of Natural History , 25:
1-26. (In Japanese, with English original descriptions.)

Medvedev, L. N. 1992. Fam. Chrysomelidae. In. Opredelitef nasekomyh Dalnego Vostoka SSSR. V. 3.
Part 2. Nauka, St. Petersburg: 533-602. (In Russian.)

Mikhailov, Y. E. and M. Hayashi, 2000. Chrysomelidae of Sakhalin I. The Entomological Review of
Japan , 55: 71-83.

(Received September 19, 2004; Accepted January 22, 2005)

Ent. Rev. Japan , 60 (1): 9-12, April 30, 2005

A New Pterostichus (Coleoptera: Carabidae)
from the Islands of Tsushima, West Japan

Seiji Morita

Higashi-gotanda 5-19-7, Shinagawa-ku, Tokyo, 141-0022 Japan

Abstract A new pterostichine carabid beetle, Pterostichus ( Micronialoe) kaniei is described
from the Islands of Tsushima, West Japan. It is related to P. ( M.) bifoveolatus Park, Kwon et
Lafer, but differs from it mainly in the body size, structure of pronotum, and shape of the
male genital organ.

So far as I am aware, only three species of the genus Pterostichus have been recorded from
the Islands of Tsushima, which are Pterostichus eschscholtzii Germer (= P. fortis Morawitz ),
P. symmetricus Straneo and P. opacipennis JedliCka. Of these, the latter two are endemic to
the islands and in this paper, I am going to describe a new species as the third endemic species.

Abbreviations. The abbreviations used herein are as follows: L—body length, measured
from apical margin of clypeus to apices of elytra; HW—greatest width of head; PW—greatest
width of pronotum; PL—length of pronotum, measured along the mid-line; PA—width of
pronotal apex; PB—width of pronotal base; EW—greatest width of elytra; EL—greatest length
of elytra; FL—length of metafemur; ML—length of metatrochanter; TL—length of hind tarsus;
M —arithmetic mean; NSMT—National Science Museum (Nat. Hist.,), Tokyo; H—holotype.

Acknowledgements. I wish to express my deep gratitude to Dr. Shun-Ichi Ueno for critical¬
ly reading the manuscript of this paper. My thanks are also due to Messrs. Noboru Kanie and
Takashi Kurihara for supplying me with important material.

Thanks are also due to Dr. German Sh. Lafer and Dr. Jong Cheol Paik for giving me an
opportunity to study the specimens belonging to the subgenus Micronialoe.

Pterostichus ( Micronialoe ) kaniei Morita, sp. nov.

(Figs. 1-7)

Description. L: 6.1-7.0 mm. Colour dark brown to blackish brown, but the appendages are
a little lighter than dorsum.

Head moderately convex; PW/HW 1.63 in H, 1.67 in 1<?, 1.64 in 1 £; frontal furrows
deep, wide and divergent posteriad, and with weak wrinkles; eyes small and less convex; lateral
grooves deep, straight and reaching the basal part of genae on each side; anterior supraorbital
pores situated a little before the mid-eye level; posterior ones situated a little behind the post¬
eye level; genae slightly convex; neck wide; microsculpture composed of polygonal meshes;

mentum tooth bifid and wide; relative lengths of antennal segments as follows:-I : II : III :

IV : V : VI : XI = 1 : 0.50 : 0.95 : 0.93 : 0.84 : 0.84 : 1.14 in H, = 1 : 0.51 : 0.92 : 0.83 : 0.75 :

10

Seiji Morita

0.76: 1.08 in 1£.

Pronotum a little convex and widest at about basal 2/5 (measured along the mid line) in H;
PW/PL 1.15 in H, 1.14 in 1 <? and 1 apex widely emarginate, a little wider than base; PA/PB
0.76 in H, 0.78 in 1 c?, 0.82 in 1?;PW/PA 1.49 in H, 1.47 in 1 d\ 1-45 in l£;PW/PB 1.13 in
H, 1.15 in 1 (?, 1.19 in 1 apical angles slightly produced and obtusely pointed at the tips, hind
ones obtuse; sides weakly and widely arcuate in front, shallowly sinuate behind, and then weak¬
ly divergent just before hind angles in H; base emarginate at middle and slightly arcuate at the
sides; median line finely impressed; anterior transverse impression evanescent, posterior one
vague; two basal foveae present on each side; inner basal fovea deep, small, linear at the bot¬
tom, and very sparsely and rather coarsely punctate at the bottom; outer basal fovea short, linear
and situated near hind angle; microsculpture composed of fine transverse meshes.

Elytra moderately convex; shoulders prominent and with a rather large tooth on each side;
EW/PW 1.27 in H, 1.26 in 1-?-; EL/EW 1.49 in H, 1.43 in 1-?-; sides slightly arcuate, and with
very shallow preapical emargination; epipleuron gradually narrowed towards apex; inner plica
distinct; apices conjointly rounded; basal border slightly arcuate and joining stria 1; intervals
weakly convex and impunctate; striae coarsely and sparsely punctate; marginal series composed
of 13 to 15 pores; microsculpture composed of fine transverse meshes.

Gula with transverse wrinkles at the sides; proepisterna, sides of prosternum, mesoster-
num, mesepisterna, and metepisterna coarsely punctate; stemites rugose; microsculpture of anal
stemite (VII) composed of polygonal to wide meshes in apical half and of wide to transverse
ones in basal half; in anal sternite elongate, weakly depressed at apical part, and narrowly
bordered throughout.

Legs stout; TL/HW 1.02 in H, 0.97 in 1 c?, 0.90 1 metafemora with two setae on each
side; metatrochanters with a seta on each side; ML/FL 0.48 in H, 0.50 in 1 d\ 0.47 in 1 £.

Aedeagus stout, small and moderately bent at basal third; viewed dorsally, apical part of
aedeagus curved towards the right and with simply rounded apex; right wall of aedeagus widely
concave, and with a carina (cf. Figs. 3-5, c) at the ventral edge; ventral side of aedeagus weakly
concave (cf. Fig. 4); apical lobe rather short and thin.

Right paramere elongate, bent at basal 3/5, and with rounded apex; left paramere wide.

Type series. Holotype: <?, Mt. Ohira-yama, 9. VI. 2002, T. Kurihara leg. (NSMT).
Paratypes: 1 <?, 1 Mt. Mokkoku-yama, 24. IX. 1993, N. Kanie leg.

Localities of the type series. Mt. Ohira-yama and Mt. Mokkoku-yama, Izuhara-machi,
Nagasaki Prefecture, Japan.

Notes. The subgenus Micronialoe is a close relative of Abea beyond all doubt. The two
subgenera have many basic characters in common, including the shape of the male genital
organ. The only prominent difference between them is the shape of the right paramere of the
male genital organ: in Abea , short and robust; in Micronialoe , elongate and bent.

The shape of the right paramere of the male genital organ serves by itself as a key charac¬
ter for a higher classification of pterostichine carabids, though there are some exceptions. I place
this new species in the subgenus Micronialoe , for the time being.

This new species may have a relationship with P. (M.) bifoveolatus Park, Kwon et Lafer
distributed in Korea. However, it is distinguished from the latter by the following points: 1)
body smaller; 2) body form robust; 3) neck narrower; 4) frontal furrows shorter and more
strongly divergent posteriad; 5) microsculpture of head more clearly impressed; 6) structure of
pronotum different (weakly produced apical angles, reflexed basal borders wider near apical

New Pterostichus from West Japan

11

Figs. 1-7. Pterostichus ( Micronialoe) kciniei Morita, sp. nov. from Mt. Mokkoku-yama.-1, Left side of

pronotum; 2, aedeagus, left lateral view; 3, aedeagus, right dorso-lateral view; 4, aedeagus, right ventro¬
lateral view; 5, aedeagus apico-dorsal view; 6, right paramere, left lateral view; 7, left paramere, left lateral
view, c - carina. (Scale 1 mm: A for 1; 0.5 mm: B for 2-7.)

angles, basal part convex and almost smooth, posterior transverse impression vague, bottom on
basal foveae shallower, and outer foveae vague); 7) elytral striae more strongly impressed; 8)
EL/EW 1.49 in <?; 9) anal sternite weakly depressed at apical part; 10) aedeagus elongate and
with rather weak carina. [in P. (M.) bifoveolatus Park, Kwon et Lafer, L: 7.7 mm. Head rather
large, with flat eyes; neck wide. Pronotum moderately convex; reflexed lateral borders very nar¬
row throughout; outer foveae deeper; PW/HW 1.58, PW/PL 1.11, PW/PA 1.43, PW/PB 1.18,
PA/PB 0.82 in 1 <?. Elytra elongate; EW/PW 1.21; EL/EW 1.65 in 1 c?; striae moderately punc¬
tate. Anal sternite in c? strongly depressed at apical part. Aedeagus robust, short and with a
prominent carina; right paramere elongate and arcuate.]

This species is named in honor of Mr. Noboru Kanie, the discoverer of the pterostichine.

12

Seiji Morita

to

1r] : -ttMMni-#** a z/(D lfffi. - a

v (7) 1 fffl, 7'>vU 'f'ffzfz, A v Pterostichus ( Micronialoe ) famin' Morita ^E2tL/:.

P. (A/.) bifoveolatus Park, Kwon et Lafer /M 3t\ fr

WITfcflflHt, tojflc-c, #ftK*B(l3*i*. *15,

Micronialoe t Abea LTtt, #< £>£>, dd-CttEBU

Ltl^SUtCL/:.

References

Morita, S., 1992. A new subgenus and species of Pterostichus (Coleoptera, Carabidae) from Aomori
Prefecture, North Japan. Elytra, Tokyo , 20: 15-19.

Park, J. K., Y. J. Kwon and G. Sh. Lafer, 1996. Classification of the genus Pterostichus Bonelli from
Korea (Coleoptera, Harpalidae) I. Micronialoe subgen. nov. Korean Journal of Entomology, Seoul,
26 : 73-77.

(Received December 16, 2004; Accepted January 22, 2005)

Ent. Rev. Japan, 60 (1): 13-16, April 30, 2005

Synonymic Notes on Two Species of the Families
Hydrophilidae and Leiodidae (Coleoptera) from Japan

Hideto Hoshina

Department of Regional Environment, Faculty of Education & Regional Studies,
Fukui University, Fukui, 910-8507 Japan

and

Masataka Sato

Dia Cuore Tokushige 306, Kamegahora 3-1404, Midori-ku, Nagoya, 458-0804 Japan

Abstract Megasternum japonicum Shatrovskiy, 1989 (Hydrophilidae) and Agathidium
(Neoceble) fujiyamaense Hoshina, 1997 (Leiodidae) are newly treated as junior synonyms of
M. gibbulum Motschulsky, 1866 and A. ( N.)funereum Angelini et De Marzo, 1990, respec¬
tively.

After the examinations of many specimens of the genera Megasternum Mulsant, 1844 of
Hydrophilidae and Agathidium Panzer, 1797 of Leiodidae upon our recent collections, and the
paratypes of Megasternum japonicum Shatrovskiy, 1989 and the holotype of Agathidium
(Neoceble) funereum Angelini et De Marzo, 1990 through the courtesies of Drs. M. Ohara
and I. Lobl, two synonymies are newly recognized as noted in the followings.

Before going further, we owe thanks to Drs. Ivan Lobl (Museum d’histoire naturelle,
Geneve) and Masahiro Ohara (The Hokkaido University Museum, Hokkaido University,
Sapporo) who kindly provided us with the opportunity to examine the type specimens.

Family Hydrophilidae

Megasternum gibbulum Motshulsky, 1866
(Figs. 1-3)

Megasternum gibbulum Motschulsky, 1866: 169; Sharp, 1884: 464; Hansen, 1999: 305.

Megastemum japonicum Shatrovskiy, 1989: 286; Shatrovskiy, 1992: 368; Hansen, 1999: 305. Syn. nov.

Distribution. Japan and Korea.

Specimens examined. Paratypes of M. japonicum , 1 ex., Mt. Kasuga, Nara Pref., Honshu, 26. V.
1954, K. Sawada leg.; 4 exs., Naga Town, Wakayama Pref., Honshu, 29. III. 1952, K. Ishikawa leg.
(Those paratypes are preserved in the collection of Systematic Entomology, Faculty of Agriculture,
Hokkaido University.); 1 ex., Mt. Eniwadake, Hokkaido, 20. VH. 1997, H. Hoshina leg.; 7 exs., Shinrin-

14

Hideto Hoshina and Masataka Sato

1 2 3

Figs. 1-3, Megastemum gibbulum Motschulsky. Median lobe of male genitalia in lateral view.
Scale: 0.5 mm.

Kagakuen, Takao, Tokyo Pref., Honshu, 22. XI. 2000, H. Hoshina leg.; 8 exs., Tendaki, Oya Town,
Hyogo Pref., Honshu, 8. VI. 1996, H. Hoshina leg.; 1 ex., Matsuyama Castle, Matsuyama City, Ehime
Pref., Shikoku, 14. VIII. 1996, H. Hoshina leg.; 1 ex., Shirataki, Nagahama Town, Ehime Pref., Shikoku,
15. VI. 1997, H. Hoshina leg.; 2 exs., Chikuzen-Okinoshima Is., Fukuoka Pref., Kyushu, 21. V. 1998, H.
Hoshina leg.; 2 exs., Shiroyama, Kagoshima City, Kagoshima Pref., Kyushu, 7. III. 1997, H. Hoshina leg.

Notes. Shatrovskiy (1989; 1992) described Megastemum japonicum on specimens from
some type specimens of Megastemum gibbulum Motschulsky in the Lewis Collection of the
Natural History Museum in London, and distinguished by the reddish brown dorsal color from
the blackish M. gibbulum Motschulsky, 1866. However, M. gibbulum are variable among
specimens upon such characters as the dorsal coloration, clearness of punctures in the striae,

Synonymic Notes of Hydrophilidae and Leiodidae

15

density of the discal punctures, strength of curvature of median lobe of aedeagus in lateral view
(Figs. 1—3), girth of median lobe in ventral view, and so on, and the character defining M.
japonicum fall evidently in the range of variation. Moreover, the specimens of both reddish
brown and black dorsal color and the two types of median lobe (Figs. 2 and 3) were collected at
the same time from one locality, Tendaki in Hyogo, and no correction between the dorsal col¬
oration and the shape of aedeagus has not been demonstrated. Consequently, it can be given as
the conclusion that M. japonicum is a junior synonym of M. gibbulum , neither independent
species nor the local variation.

Family Leiodidae

Agathidium {Neoceble) funereum Angelini et De Marzo, 1990

Agathidium ( Neoceble) funereum Angelini et De Marzo, 1990: 92; Angelini, 1995: 140; Hoshina, 2000: 67.
Agathidium (Neoceble) fujiyamaense Hoshina, 1997: 164. Syn. nov.

Distribution. Japan (Honshu).

Specimens examined. Holotype of A. (N.) funereum, c?, Yaseyuen, Kyoto Pref., Honshu, 4. VIII.
1980, C. Besuchet leg. (preserved in Museum d’histoire naturelle, Geneve); holotype of A. (N.) fujiya¬
maense , d\ Aokigahara, Mt. Fujisan, Yamanashi Pref., Honshu, 23. VIII. 1982, S. Naomi leg. (preserved
in Kyushu University). Not type specimens, 5 exs., Tendaki, Oya Town, Hyogo Pref., 28. V. 1998, H.
Hoshina leg.; 16 exs., Mt. Maya, Kobe City, Hyogo Pref., 30. V. 1998, H. Hoshina leg.

Notes. Agathidium (Neoceble) fujiyamaense was described on a male from Aokigahara,
Mt. Fujisan, and A. (N.) funereum on the specimens from Kyoto. The median lobe of aedeagus
in A. (N.) funereum is a little slenderer than that in A. (N.) fujiyamaense in ventral view (cf. figs
in Angelini and De Marzo, 1990, and Hoshina, 1997), but these differences seem to be not
enough for separating both populations as the independent species nor subspecies after exami¬
nation of some additional specimens.

im • \m IE# : -

Megasternum japonicum li Shatrovskiy (1989) Cio T, M. gibbulum

Motschulsky, 1866 : •feYjl/Y^v^y) <T>9 d

b . M. gibbulum Motschulsky, 1866 i ±tMM

& & <D "C (i & V L < , M. japonicum (i b l- M. gibbulum

b. itiy Agathidium (Neoceble) fujiyamaense Hoshina, 1997
n A v) (±, S3cii#^ ? A. (N.) funereum Angelini et De Marzo, 1990
7^7 nAy) oodrtl t < , Cl b (D<D t 3Eft * fiU

t L tz.

16

Hideto Hoshina and MasatakaSATO

References

Angelini, F., 1995. Revisione tassonomica delle specie paleartiche del genere Agathidium Panzer
(Coleoptera: Leiodidae: Agathidiini). Museo Regioncile di Scienze Naturali, Torino, Monografie , 18:
1-485.

-and L. De Marzo, 1990. Anisotomini del Giappone (Coleoptera, Leiodidae). Entomologica,

Bari , 23: 47-122.

Hansen, M., 1999. Hydrophiloidea (s. str.) (Coleoptera). World Catalogue of Insects, 2. 416 pp. Apollo
Books, Stenstrup.

Hoshina, H., 1997. Two new species on the genus Agathidium (Coleoptera, Leiodidae) from Japan. The
Japanese Journal of Systematic Entomology , 3: 161-165.

-, 2000. A taxonomic study on the subgenus Neoceble (Coleoptera: Leiodidae: Agathidium) from

Kyushu, Japan. Species Diversity, Sapporo. 5: 59-88.

Motschulsky, V., 1866. Catalogue des Insects re$us du Japon. Bulletin de la Societe imperiale des
Naturalistes de Moscou , 39,1: 163-200.

Sharp, D., 1884. The water-beetles of Japan. Transactions of the Entomological Society of London, 1884:
439^164.

Shatrovskiy, A. G., 1989. Hydraenidae, Hydrophilidae (pp. 260-293). In Ler, P. A. (ed.). Operedelitel’
nasekomykh Dal’nego Vostoka SSSR v shesti tomakh. Vol. 3. Zhestkokrylye, ili zhuki (part 1). 572
pp. Nauka, Leningrad. (In Russian.)

Shatrovskiy, A. G., 1992. Novye i moloizvestnye vodolyubovye (Coleoptera, Hydrophiloidea) iz
yuzhnogo Primor’ya i sopredel’iykh territorii. Entomologicheskoe Obozrenie , 71: 359-371. (In
Russian.)

(Received December 16, 2004; Accepted January 28. 2005)

Ent. Rev. Japan , 60 (I): 17-21, April 30, 2005

The Complex of Trechiama fujitai (Coleoptera: Trechinae)
from Hyogo Prefecture, West Japan (III)

— A New Relative of Trechiama latilobatus Ashida —

Hisashi Ashida

7-4-201, Shimeien, Ibaraki, Osaka, 567-0045 Japan
E-mail: BYD01621@nifty.ne.jp

Abstract An additional new species belonging to the Trechiama fujitai complex in the group
of T. oni is described from Asago-cho and Aogaki-cho in the central part of Hyogo Prefecture
under the name T. (s. str.) asagonis Ashida, sp. nov. This new species is a close relative of T.
(s. str.) latilobatus Ashida, though easily discriminated from the latter species by the configu¬
ration of the male genitalia.

In the first part of this series, I described Trechiama latilobatus from the central part of
Hyogo Prefecture, which is an isolated species within the fujitai complex of the group of T. oni
(Ashida, 2003). This species was found from three localities in the eastern hills of the upstream
of the Maruyama-gawa River that flows from the central part of the prefecture to the Sea of
Japan. This is quite exceptional in the fujitai complex, because this complex is mainly distrib¬
uted in the western area of the Maruyama-gawa / Ichi-kawa line and another complex of T.
kosugei largely occupies the opposite side of the line (Ueno, 1985; Ashida, 2003, 2005). When
I described T. latilobatus , I also examined several specimens brought from the adjacent areas of
its localities; they are similar to that species but have some differences. However, I did not
determine the real status of them due to the insufficient number of available specimens. After
that, I confirmed the stable phenotype of this population by examining a series of specimens,
and am convinced that this is a new species.

The abbreviations used herein are as follows: HW - greatest width of head; PW - greatest
width of pronotum; PL - length of pronotum, measured along the mid-line; PA - width of
pronotal apex; PB - width of pronotal base; EW - greatest width of elytra; EL - greatest length
of elytra; M - arithmetic mean. Measurement was carried out using six male and six female
specimens.

Trechiama (s. str.) asagonis Ashida, sp. nov.

(Figs. 1-5)

Length: 4.75-5.45 mm (from apical margin of clypeus to apices of elytra).

Closely related to T. latilobatus Ashida (2003, p. 431, figs. 1, 4-7), though discriminated
from the latter species by less depressed and less elongate shape of male genitalia.

18

Hisashi Ashida

Fig. 1. Trechiama (s. str.) asagonis Ashida, sp. nov., from the Iyu-toge in Asago-cho, <f, dorsal view.

Relatively small species and externally very similar to T. latilobatus. Color yellowish
brown with light-colored appendages. Head slender, a little longer than width; genae slightly
convex; antennae slender, reaching the middle of elytra. Pronotum similar to that of T. latiloba¬
tus though a little wider, subcordate, widest at two-thirds from base; PW/HW 1.37-1.46 (M
1.42), PW/PL 1.06-1.18 (M 1.11), PW/PA 1.31-1.47 (M 1.41), PW/PB 1.27-1.45 (M 1.39),
PB/PA 0.97-1.06 (M 1.01); sides more strongly arcuate in front; front and hind angles as in T.
latilobatus ; postangular setae absent; base slightly emarginated at middle. Elytra exactly similar
to those of T. latilobatus , regularly ovate, rather slender, widest at about middle; EW/PW
1.58-1.76 (M 1.69); EL/PL 2.68-2.94 (M 2.78); EL/EW 1.42-1.52 (M 1.48); EW/PW
1-67—1.82 (M 1.73); EL/PL 2.50—2.75 (M 2.67); EL/EW 1.45—1.51 (M 1.49); prehumeral bor¬
ders oblique; shoulders completely effaced; sides regularly rounded towards apices; striae and
chaetotaxy as in T. latilobatus. Legs as in T. latilobatus.

Male genital organ large, elongate and heavily sclerotized, basically similar to that of T.
latilobatus. Aedeagus one-third as long as elytra, compressed on dorsum though less heavily
than in T. latilobatus ; basal part ampler and more strongly curved ventrad than in T. latilobatus
with large and hyaline sagittal aileron; viewed laterally, middle part moderately convex on dor¬
sum, then gradually and regularly narrowed towards apical tip, which is much thicker than in T.
latilobatus ; viewed dorsally, apical lobe gradually dilated, widest at apical third, and then feebly

Trechiamci fujitai Complex from Hyogo Prefecture (III)

19

-1

0.5 mm

Figs. 2-5. Male genitalia of Trechiama (s. str.) asagonis Ashida, sp. nov., from the Iyu-toge in
Asago-cho; left lateral view (2), dorsal view (3), and separated copulatory piece, lateral (4)
and dorsal (5) views.

narrowed apicad, which is wide and subsquare with widely rounded corners; viewed ventrally,
apical lobe longitudinally convex behind apex. Inner sac armed with a teeth-patch, a copulatory
piece and two plates as in T. latilobatus ; teeth-patch small, consisting of fairly long teeth, lying
on left side at about middle of aedeagus; copulatory piece very lightly sclerotized, lying at right
side of teeth-patch, trapezoidal, one-ninth as long as aedeagus, a little wider than in T. latiloba¬
tus , rolled ventrad, whose apical margin is slightly emarginated and projected at right-apical
corner; two large plates covered with minute scales at the dorsal side of apical orifice. Styles
nearly straight and long; the left one slightly longer than the right, each bearing three or four
setae at apex.

Type series. Holotype: <?, 23. IX. 2003, H. Ashida leg. Paratypes: 1 <T, 1 *, 1. VI. 2002,
A. Souma leg.; 3<?<T, 4?*, 14. IX. 2002, A. Souma leg.; 3 d* 1-?-, 4. V. 2003, S.
Yamashita leg.; 2. VIII. 2003, Y. Okuda and T. Saito leg.; 13dV, 12* *, 23. IX.

2003, H. Ashida leg.; 4<?<T, 1*, 5. X. 2003, Y. Okuda leg. The holotype and one female
paratype are preserved in the collection of the National Science Museum (Nat. Hist.), Tokyo.

20

Hisashi Ashida

Type locality. Iyu-toge (400 m in altitude), the southeastern slope of Mt. Asago-yama
(756.5 m in height), Kawakami, Asago-cho, Hyogo Prefecture, West Japan. Asago-cho was
incorporated into Asago-shi on April 1, 2005.

Further records. 1 cT, Mt. Awaga-yama (700 m in altitude), Aogaki-cho, Hyogo
Prefecture, 13. VII. 2001, M. Mori leg.; 1 d\ 1?, same locality (550 m in altitude), 31. III.
2002, M. Mori leg.; 1-?-, same locality (700 m in altitude), 21. IV. 2002, Y. Okuda leg.; I?,
same locality (700 m in altitude), 2. V. 2002, T. Saito leg. Aogaki-cho was incorporated into
Tamba-shi on November 1, 2004.

Etymology. The specific name of this species is derived from the type locality, Mt. Asago-
yama in Asago-cho.

Notes. As described above, T. asagonis is a close relative of T. latilobatus, which is one
of the most isolated and specialized species within the fujitai complex. Judging from the fea¬
tures of the male genitalia, however, the former species is less specialized than the latter, name¬
ly, the aedeagus of T. asagonis is less flattened, less elongated at apical part, and somewhat
resembles those of the other members of the fujitai complex. The Iyu-toge, the type locality of
the present species, is located on the southeastern slope of Mt. Asago-yama and is 3.5 km dis¬
tant to the northeast from Tataragi, the type locality of T. latilobatus. The type specimens were
dug out from colluvia deposited at the riverhead of the lyudani-gawa River, an eastern branch of
the upstream of the Maruyama-gawa River. The second locality of this species lies on the east¬
ern slope of Mt. Awaga-yama (962.3 m in height). It is 6.5 km distant to the east-northeast from
the Iyu-toge, 7 km to the northeast from Kurogawa, the second known locality of T. latilobatus ,
and 2.5 km to the south from the Tosaka-toge, the type locality of T. siva Ashida, a member of
the notoi complex. Trechiama asagonis from Mt. Awaga-yama is therefore the easternmost
population of the fujitai complex. Two collecting points on Mt. Awaga-yama are near the head
of the Inazuchi-gawa River, one of the sources of the Kako-gawa River that flows into the Seto
Inland Sea. There, T. asagonis coexists with another species belonging to the kosugei complex
that is dominant. Details will be reported elsewhere.

Acknowledgements

I thank Messrs. Masato Mori, Akinao Souma, Shun-ichi Yamashita, Yoshihide Okuda,
and Takumi Saito for their kindly providing the materials, and Dr. Shun-ichi Ueno of the
National Science Museum (Nat. Hist.), Tokyo, for his continuous guidance.

PS X : ^ ? y A i/fh (Ulllfl*) — 9 * 9 * y \£ rf 5 A y

1 fffl . - 9 9 y 9 v kf rf ^ A y Trechiama (s. str.) latilobatus

Ashida, 2003 (i, CKCffil) frbtZWLZtitz, 7y^^7f

t btifzW, T-fl-nV ? yf-tfrf

* A v T. (s. str.) asagonis Ashida, sp. nov. L, Efc L tz. 9 9 y ¥ *

Trechiama fujitai Complex from Hyogo Prefecture (III)

21

References

Ashida, H., 2003. The complex of Trechiama fujitai (Coleoptera, Trechinae) from Hyogo Prefecture,
West Japan (I) —Two new species from the Maruyama-gawa drainage area—. Elytra , Tokyo , 31:
431-438.

-, 2004. An additional species belonging to the Trechiama notoi complex (Coleoptera, Trechinae)

from the southern part of the Tajima area in Hyogo Prefecture, Central Japan. Elytra , Tokyo , 32:
259-263.

-, 2005. The complex of Trechiama fujitai (Coleoptera, Trechinae) from Hyogo Prefecture, West

Japan (II) —Two new species and several new records from the Ibo-gawa drainage area—. Elytra ,
Tokyo , 33: in press.

Ueno, S.-I., 1985. The group of Trechiama oni (Coleoptera, Trechinae) —Its distribution and differentia¬
tion—. Memoirs of the National Science Museum, Tokyo, (18): 163-198.

(Received January 25, 2005; Accepted February 16, 2005)

Ent. Rev. Japan , 60 (1): 23-33, April 30, 2005

Origin of Ohomopterus uenoi (Coleoptera: Carabidae: Carabinae) as
Deduced from Comparisons of DNA Sequences of Mitochondrial
ND5 Gene and Nuclear Internal Transcribed Spacer I (ITS I) with

Morphological Characters

Osamu Tominaga”, Yuki Imura 2 ’, Munehiro Okamoto”, Zhi-Hui Su 4 * 1 ,
Tooru Ojika 5) , Nobuo Kashiwai 61 and Syozo Osawa 71

1) A312,4-1 -15, Shibatsuji-cho, Nara, 630-8 ] 14 Japan

2) Shinohara-cho 1249-8, Kohoku-ku, Yokohama, 222-0026 Japan

3) Department of Laboratory Animal Science, School of Veterinary Medicine,
Faculty of Agriculture, Tottori University, Tottori, 680-8553 Japan

4) JT Biohistory Research Hall, 1-1 Murasaki-cho, Takatsuki, Osaka, 569-1125 Japan

5) Imahon-Machi, 2-9-27-301, Anjo-shi, Aichi, 446-0008 Japan

6) Hosen-Gakuen High School, Chuo, Nakano-ku, Tokyo, 164-8628 Japan

7) Ushita-Asahi 2-4-7-1003, Higashi-ku, Hiroshima, 732-0067 Japan

Abstract The phylogenetic trees have been constructed using the mitochondrial ND5 gene
and the nuclear ITS I sequences of Ohomopterus uenoi from the Kongo Mountains together
with its relevant species inhabiting the Kinki and Chubu Districts in Central Japan. The mor¬
phological characters, especially the genital organs of these species, have been re-examined.
Altogether, the results obtained from molecular and morphological characteristics suggest
strongly that O. uenoi is a descendant from a hybrid between the female of O. arrowianus
and the male of either O. iwawakianus or O. kiiensis.

Introduction

Ohomopterus uenoi (Ishikawa, 1960) is one of the most peculiar species of the genus
Ohomopterus in several respects. Firstly, the distribution of this species is narrowly restricted to
the upper parts of Mt. Kongo-zan and Mt. Yamatokatsuragi-san of the Kongo Mountains in the
Kinki District of central Honshu, Central Japan (Ishikawa, 1962; Hiura, 1965; Komiya, 1971;
Kinki Research Group of Carabid Beetles, 1979, Ishikawa, 1991, etc.). No species other than
O. uenoi inhabiting such an isolated region have been known in the genus Ohomopterus.
Secondly, O. uenoi has an exceptionally huge copulatory piece of the male genital organ of the
insulicola-typc (I-type). Thirdly, except for the copulatory piece, the general appearance of O.
uenoi is similar to O. yaconinus and to some extent to O. iwawakianus both having copulatory
piece of the yaconinus-type (Y-type) and inhabiting with O. uenoi in the Kongo Mountains,
although the latter two species are distributed widely in the Kinki District. Ohomopterus
dehaanii and O. yamato also inhabit the Kongo Mountains, both of which would most probably

Corresponding author. E-mail: su.zhihui@brh.co.jp

24

Osamu Tominaga et al.

be irrelevant to O. uenoi and are not treated here.

Now, where did O. uenoi come from? In one of our previous papers (Su et al. , 1996), we
reported that the mitochondrial ND5 gene of O. uenoi shares the common ancestry with that of
O. arrowianus , the main distributional range of which is the central Honshu. There are at least
two possibilities to account for the origin of O. uenoi. One would be that there was a certain
period where the ancestor of O. arrowianus was distributed in Central Japan through the Kongo
Mountains. The male copulatory piece of O. arrowianus evolved to the direction of extreme
elongation. Another possibility would be that the ancestor of O. arrowianus (-?-), which once
inhabited the Kongo Mountains, was hybridized with some other Ohomopterus species (d*)
having inhabited there. This would have resulted in the ancestor of the present-day O. uenoi.
Ohomopterus arrowianus that participated in this hybridization must be the female, because the
mitochondrial inheritance is female mediated, and the ND5 gene of O. uenoi reveals the O.
arrowianus- type sequence.

To know which possibility is more likely than the other one, we have analyzed the
sequences of the ND5 gene and nuclear ITS I of all the Ohomopterus species and subspecies
inhabiting the Kinki and Chubu Districts including O. uenoi , O. arrowianus, O. iwawakianus
and O. yaconinus. The whole pictures of the phylogenetic trees will be published later.

If the ITS I sequence of O. uenoi is of the O. arrowianus- type, the first possibility, i.e., the
hypertrophic evolution (probably by genetic event(s)) of the male genital organ of O.
arrowianus to the O. uenoi- type would be the case. If the ITS I sequence is of some other
species, the second possibility, i.e., O. uenoi would have been derived from the hybrid ancestor
between O. arrowianus (-¥-) and the species concerned (<?).

Materials and Methods

More than 150 Ohomopterus specimens from the Kinki and Chubu Districts were ana¬
lyzed, and only those relevant to O. uenoi were treated in this paper and are shown in the phylo¬
genetic trees. The ND5 gene sequence (1,069 bp.) analysis and construction of a phylogenetic
tree by the UPGMA with bootstrap test were the same as described previously (e.g., Su et al .,
2004). The dating was done assuming that a 0.01 unit corresponds to 3.6 million years for the
carabid ND5 gene according to Su et al. (1998, 2001).

The DNA fragment including ITS I, 5.8S rDNA and ITS2 was amplified by two adapted
primers: 5’-AAG TCG TAA CAA GGT TTC CG- 3’ and 5’-TCC TTG TTA GTT TCT TTT
CCT C-3\ About 1,000 bp sequence region of ITS I was directly sequenced whenever possible
and used for construction of the phylogenetic tree. The direct sequencing was sometimes not
possible by a small heterogeneity due to multicopy of ITS I. In such cases, the amplified DNA
fragments were subjected to cloning. Five to ten independent clones from one individual were
sequenced.

For both the ND5 gene and ITS I, phylogenetic analyses by means of the UPGMA, NJ-,
MP- and ML-methods (see Saito et al., 2003) were performed, obtaining essentially the same
results. Only the UPGMA trees were shown in this paper. Both the ND5 and ITS I trees are also
used in the next paper (Imura et al., 2005a).

Origin of Ohomopterus uenoi

25

Results

Phylogenetic trees of the representative Ohomopterus species and subspecies, which are
supposed to be relevant to O. uenoi

ND5 tree: Fig. 1 shows the UPGMA-phylogenetic tree of the representative Ohomopterus
species and subspecies in the Kinki and Chubu Districts with exclusion of O. yamato , O. insuli-
cola and O. albrechti which are not directly connected with the present subject. Four major
clusters were recognized in the tree. The cluster I consisted of mostly O. arrowianus from the

1 -maiyasanus (Azai. Shiga)

2- arrowianus nakamurai (Hase. Nagano)

3 - insulicola (Fujimi, Nagano)

4 - insulicola (Otari, Nagano)

5 - insulicola (Sado, Niigata)

6 - insulicola (Sado, Niigata)

7- arrowianus (Nagoya, Aichi)

8- arrowianus (Toyota, Aichi)

9- insulicola (Kisofukushima, Nagano)

I O-maiyasan us (N agara. G i fu)

II -arrowianus (Neba, Nagano)

12 - arrowianus (Toyota, Aichi)

13 - arrowianus (Asuke. Aichi)

1 4- arrowiamis (Hino, Gifu)

15 - arrowianus (Nagara, Gifu)

16 - arrowianus (Hinokita, Gifu)

17 - arrowianus (Okazaki. Aichi)

18 - arrowianus (Toyohashi, Aichi)
)9-maiyasanus (Itoigawa, Niigata)

20 -yaconinus blairi (Wajima. Ishikawa)

21 -yaconinus blairi (Nanao. Ishikawa)

22 - maiyasanus (Shirotori. Gifu)

23- arrowianus (Nagiso, Nagano)

24- arrowianus (Nagiso, Nagano)

25 - maiyasanus X iwawakianus (Suzuka, Mie)

26- maiyasanus suzukanus (Komono. Mie)

27 - maiyasanus suzukanus (Yokkaichi, Mie)
2%-arrowianus kirimurai (Mihama, Mie)

29- arrowianus kirimurai (Mihama. Mie)

30- tnaiyasanus takiharensis (Odai. Mie)

31 -maiyasanus (Shirotori. Gifu)

32 - arrowianus komiyai (Haruno, Shizuoka)

33- arrowianus komiyai (Kakegawa. Shizuoka)

34 - uenoi (Gose, Nara)

35- uenoi (Gose. Nara)

36- uenoi (Gose. Nara)

37- uenoi (Gose, Nara)

38 - uenoi (Mt. Kongo, Nara)

39 - maiyasanus (Kanzaki, Hyogo)

40- dehaanii (Sayoh, Hyogo)

4\-maiyasanus (Takefu, Fukui)

42 -maiyasanus (Sueno, Fukui)

| Cluster IV

99

Fig. 1. UPGMA-phylogenetic tree of the mitochondrial ND5 gene of the representative Ohomopterus species
and subspecies of the Kinki and Chubu Districts. The tree is based on that presented by Su et al. (1996),
with additions of the sequences determined in this study. All examples of O. insulicola, O. maiyasanus ,
O. yaconinus and O. dehaanii in the cluster I are the hybrid- or hybrid-derivatives (to be published else¬
where). For the clusters II-IV, see the first section of Results. Scale bar shown in all the trees in this
paper represents Kimura's 2-parameter evolutionary distance. Value at the node shows bootstrap confi¬
dence level (%) based on 500 resampling. For details, see Su et al. (2004).

26

Osamu Tominaga et al.

(a)

Cluster A

for details, sec Fig. 2b

0.002

| Cluster (5]

32-arrowianus komiyai (Haruno. Shizuoka)
\09b-moiyasanus (Shirakawa, Gifu)

23 -arrowianus (Nagiso, Nagano)

I \l-maiyasanus (Aogaki. Hyogo)

1 \6-maiyasanus (Omiya, Kyoto)

1 9-maiyasanus (Itoigawa. Niigata)

46 -arrowianus murakii (Toba. Mie)
arrowianus murakii (Toba, Mie)

43 -arrowianus murakii (Futami, Mie)
50-arrowianus murakii (Seivva, Mie)
2-arrowianus nakamurai (Hase, Nagano)

1 Ob-maiyasanus (Nagara, Gifu)

1 \2-maiyasanus (Wajima, Ishikawa)

28 -arrowianus kirimurai (Mihama, Mie)
14b -arrowiamus (Hino. Gifu)

1 1-arrowianus (Neba, Nagano)
insulicola (Minowa, Nagano)

9 -insulicola (Kisofukushima. Nagano)
inxulicola (Sado, Niigata)

68b -maiyasanus (Ibaraki. Osaka)
39-maiyasanus (Kanzaki. Hyogo)
maiyasanus (Sekinontiya. Hyogo)

60 -maiyasanus (Nanko, Hyogo)

115 -maiyasanus (Yoka. Hyogo)

japonicus-Yineage

dehaanii -\ineage

insulicola-\\ neage

yaconinus -lineage

albrech ti/yamato -lineage

Fig. 2. UPGMA-phylogenetic tree of the nuclear ITS I sequence of the representative Ohomopterus species
and subspecies of the Japanese Islands with special reference to the Kinki and Chubu Districts, a) the
overall tree; b) the details for Cluster A. Some bootstrap values are shown at the branching points. All
examples of O. yaconinus, O. maiyasanus and O. arrowianus in the cluster A are the hybrid- or hybrid-
derivatives (to be published elsewhere). All examples of O. insulicola in the cluster B are the hybrid- or
hybrid-derivatives between O. arrowianus (£) and O. insulicola (cf). The specimens without number
represent the lack of the ND5 sequences.

Chubu District with some individuals presumably derived from hybrids between O. arrowianus
(£) and other species (<?). Five individuals of O. uenoi collected both from Mt. Kongo-zan and
Mt. Yamatokatsuragi-san of the Kongo Mountains were also included in this cluster as previ¬
ously reported (Su et al., 1996). The cluster II contained O. maiyasanus from the northern parts
of the Kinki District to Ishikawa Prefecture of the Hokuriku District along with the coast of the
Sea of Japan. The composition of the cluster III was quite complex, including O. arrowianus
murakii (!), several subspecies of O. maiyasanus, O. yaconinus, O. iwawakianus and O.
dehaanii. The cluster IV was mainly composed of O. kiiensis**. Some O. maiyasanus sub¬
species were also included in this cluster. From the results, it is apparent that O. uenoi is some¬
how related to O. arrowianus and no evidence was found the relation between O. uenoi and O.
yaconinus.

** In the present paper, the taxon “ kiiensis ” is applied to the name for a full species, which has previously been treated
as a subspecies of O. iwawakianus . Instead, O. iwawakianus iwawakianus, which is conventionally called here O.
iwawakianus, is a hybrid-derivative between O. kiiensis and O. maiyasanus, and would be, strictly speaking, not a good
species (see Discussion and Imura et al, 2005b).

Origin of Ohomopterus uenoi

27

14- arrowianus (Hino, Gifu)

98 -iwawakianus narukawai (Hisai, Mie)
%5-iwawakianus (Mt. Kongo, Nara)

68 -maiyasanus (Ibaraki, Osaka)

128- iwawakianus kiiensis (Kozagawa, Wakayama)
125 -iwawakianus kiiensis (Shingu. Wakayama)

121 -iwawakianus tmtro (Nabari, Mie)

86 -yaconinus (Gose, Nara)

1 39-maiyasanus (Hokusei, Mie)

1 32-hvawakianus kiiensis (Miyagawa, Mie)

122 - iwawakianus kiiensis (Higashiyoshino. Nara)
19-iwawakianus (Mt. Kongo. Osaka)

59- iwawakianus (Asuka, Nara)

45 -arrowianus murakii (Futami, Mie)

54- iwawakianus (Kameyama. Mie)

1 34-iwawakianus kiiensis (Kumano. Mie)

41 -maiyasanus (Takefu, Fukui)

26 -maiyasanus suzukanus (Komono, Mie)

71 -maiyasanusyoroensis (Hokusei. Mie)

\4\-maiyasanus suzukanus (Suzuka. Mie)

14-iwawakianus (Nara. Nara)
iwawakianus narukawai (Hisai. Mie)

10-maiyasanus shigaraki (Konan, Shiga)

81 -yaconinus (Matsuzaka, Mie)

138 -maiyasanus suzukanus (Komono, Mie)

62- yaconinus (Fujiwara, Mie)

64- iwawakianus shima (Takihara, Mie)

48 -maiyasanus ohkawai (Ago, Mie)

51 -iwawakianus shima (Toba, Mie)

41-iwawakianus shima (Toba, Mie)

123- iwawakianus kiiensis (Kiinagashima. Mie)

63 - maiyasanus takiharensis (Miyagawa, Mie)

38 -uenoi (Mt. Kongo. Nara)

31-uenoi (Gose. Nara)

34- uenoi (Gose, Nara)

114 -maiyasanus (Asahi, Toyama)

109 -maiyasanus (Shirakawa, Gifu)

3]-maiyasanus (Shirolori, Gifu)

13-maiyasanus (Nishiazai, Shiga)

119 -maiyasanus (Ono, Fukui)

22 -maiyasanus (Shirotori. Gifu)

\0-maiyasanus (Nagara. Gifu)

ITS I tree: Seven clusters (clusters A~G) were recognized in the ITS I phylogenetic tree
in the genus Ohomopterus from Japan (Fig. 2a), and four of them (clusters A, B, D and F)
which could be related to O. uenoi, were treated in details in this paper. The composition of the
cluster A was complex and of special importance in unmasking the origin of O. uenoi***. This
cluster contained O. iwawakianus, O. kiiensis , several subspecies of O. maiyasanus and O.
uenoi. The ITS from all the specimens of O. uenoi, O. kiiensis and O. iwawakianus fell under
the cluster A (Fig. 2b). The cluster B included both O. arrowianus arrowianus, O. a. murakii
and several subspecies of O. maiyasanus (Fig. 2a). However, the sequence difference between
them was so small that the separation was not possible in the UPGMA-tree, although O.
arrowianus and O. maiyasanus were somewhat indefinitely separated from each other in the NJ-
tree (not shown). So we tentatively treated them collectively as the members of the same cluster
B. The cluster D consisted of exclusively O. dehaanii, the individuals from the western Japan
being also included in this cluster, in contrast to the fact that in the phylogenetic tree of the ND5
gene of O. dehaanii of the Kinki and Chubu Districts were not clustered with the same species
from the western Japan (Su et al., 1996). This will be discussed in a separate paper. The cluster

*** Ohomopterus uenoi was previously considered as a derivative of a hybrid between O. arrowianus and O. yaconinus
(see Table 7.2 on p. 116 of Osawa et al ., 2004). This is not correct as discussed in this paper, because the ITS I of O.
yaconinus does not belong to the cluster A. The specimens of “ yaconinus ” (Nos. 62, 81 and 86 in Fig. 2) have been
shown to be the hybrids formed by complex participation of O. yaconinus and O. iwawakinaus (Okamoto et al., 2005).

28

Osamu Tominaga et al.

Fig. 3. Genital organ of Ohomopterus uenoi (Mt. Yamatokatsuragi-san, Osaka~Nara Prefs.). a, aedeagus in
right lateral view; b, apical part of aedeagus in the same view; c, ditto in dorsal view; d, fully everted
endophallus in right lateral view; e, ditto in posterior view; f, ditto in anterior view; g, copulatory piece
in ventral view; h, ditto in lateral view; i, peripheral rim of gonopore in right lateral view; j and k, inner
plate of vaginal apophysis in dorsal view. Scale: 2 mm for a, d-f; 1 mm for b, c, g-i, 0.5 mm for j-k.

F was composed of solely O. yaconinus. As in the case of O. dehaanii , the Kinki population of
O. yaconinus was clustered together with O. yaconinus from the same species from western
Japan, although in the ND5 tree the Kinki population was not clustered with O. yaconinus from
the western Japan (Su et al ., 1996; Okamoto et al 2005).

Morphology

Male and female genital organs of O. uenoi and its relevant species were illustrated in figs.

3-7.

The aedeagus of O. uenoi is very large and stout, nearly three-fifths of elytra in length,
with the basal portion thick, robust and twisted from the left proximal to the right apical. The
median portion is moderately arcuate in the lateral view and rather sigmoid in the dorsal view.
The apical portion strongly bent ventrally and gradually narrowed toward the apex, which is not
so sharply pointed. The endophallus is also long and tube-like, entirely lacking the basal lobes,
any inflation in the median portion and the protuberance of the praeputial pad, but only para-
ligula is well developed. The copulatory piece is monstrously elongated, about seven-eighths as
long as the aedeagus, extending rectangularly to the right side of the endophallus, acutely
curved and twisted at the base, feebly arcuate and rather compressed in the median portion,
somewhat tuberculate on the dorsal margin before the apex and gently bent near the apex which

Origin of Ohomopterus uenoi

29

Fig. 4. Genital organ of Ohomopterus arrowianus (Toyota-shi, Aichi Pref.). Captions for each illustration
correspond to those in fig. 3. Scale: 2 mm for a, d-f; 1 mm for b, c, g-i; 0.68 mm for j.

is narrowly rounded. The peripheral rim of gonopore is well sclerotized and pigmented. All
these findings are quite exceptional for the genus, and it is difficult to elucidate the true affinity
of O. uenoi by the male genitalic morphology alone, except that its copulatory piece is of I-type
and not of Y-type as in O. yaconinus or O. iwawakianus.

The inner plate of the vaginal apophysis is subquadrate in the shape and not deeply con¬
cave, resembling that of O. iwawakianus but is much different from those of O. yaconinus or O.
arrowianus.

It has been often said that the external appearance of O. uenoi resembles that of O. yaconi¬
nus , and to some extent that of O. iwawakianus both also inhabiting the Kongo Mountains.
However, O. uenoi has some features suggesting its affinity to O. arrowianus than to O. yaconi¬
nus , for example, not remarkably punctate lateral parts of the abdominal sternites, apparently
angulate inner margin of the male protibia, etc. These characteristics have not necessarily been
taken into consideration for a long time, simply because O. arrowianus is not distributed near
the Kongo Mountains. However, this possibility should be revived because, as noted before (Su
et al , 1996, and also see below), the ND5 sequence of O. uenoi is of the O. arrowianus- type
and is not the O. yaconinus- type.

In the Ohomopterus taxonomy, the copulatory piece of the male genital organ has been
pinpointed more seriously than the external morphology, because morphology of the copulatory
pieces is characteristic to the species or the species-group, while external morphology reveals
considerable similarity between various Ohomopterus species and yet sometimes conspicuous
variability even within the species.

30

Osamu Tominaga et cil.

Fig. 5. Genital organ of Ohomopterus yaconinus (<?, Gose-shi, Nara Pref.; Hisai-shi, Mie Pref. (j) and
Ureshino-cho, Mie Pref.). Captions for each illustration correspond to those in fig. 3. Scale: 2 mm for
a, d-f; 1 mm for b, c, g-i; 0.6 mm for j-k.

Discussion

In agreement with the result reported by Su et al. (1996), the ND5 gene of all examples of
O. uenoi examined in this study is included in the cluster I, to which authentic O. arrowianus
belongs. This suggests that mitochondrial DNA of O. uenoi was derived from O. arrowianus. In
contrast to the ND5 gene, the ITS I sequence of O. uenoi belongs to the cluster A, in which O.
iwawakianus, O. kiiensis and some subspecies of O. maiyasanus are included, and is not in the
cluster B where ITS I from all O. arrowianus species are included. Thus, incongruence between
the ND5 gene and ITS I of O. uenoi suggests strongly that the first possibility described in
Introduction, i.e., O. uenoi evolved from O. arrowianus to the direction to develop a huge copu-
latory piece, may be excluded. The existence of the ITS I sequence of O. uenoi in the cluster A
may be explained in such a way that O. uenoi was derived from a hybrid between O. arrowi¬
anus (?) and one of the members in the cluster (<?). The most plausible candidate would be O.
iwawakianus ( S ') that participated in the hybridization with O. arrowianus (?), although partic¬
ipation of O. kiiensis, instead of O. iwawakianus, in the hybridization cannot be excluded.
Indeed, O. iwawakianus inhabits around the area of the Kongo Mountains, and female copulato-
ry organ of O. iwawakianus reveals a strong resemblance to that of O. uenoi as compared with
that of the other species in the cluster A such as several subspecies of O. maiyasanus. It should
be noted here that O. iwawakianus is not itself an authentic species strain, and is a hybrid-deriv¬
ative between O. kiiensis (cT) and O. maiyasanus (?), i.e., mitochondria of all O. iwawakianus
were derived from O. maiyasanus, while O. kiiensis had not passed hybridization since its estab¬
lishment, i.e., it is a pure species strain (Imura et al., 2005b). Incidentally, the subspecies of O.
maiyasanus in the ITS cluster A carry the ITS sequences resembling that of O. iwawakianus.

Origin of Ohomopterus uenoi

31

Fig. 6. Genital organ of Ohomopterus iwawakianus (<?, Mt. Kongo-zan, Osaka Pref.; Matsuzaka-shi,
Mie Pref.). Captions for each illustration correspond to those in fig. 3. Scale: 2 mm for a, d-f; 1 mm
for b, c, g-i; 0.64 mm for j.

suggesting that these subspecies were derived from the hybrids between O. maiyasanus and O.
iwawakianus or O. kiiensis (Imura et al., 2005b).

From the above discussion, we suggest that most of the <9. arrowianus population, which
once inhabited between Central Japan and the Kongo Mountains, became extinct. Then the pop¬
ulation of the Kongo Mountains was geographically isolated, and hybridized with <9. iwawaki¬
anus , followed by the hypertrophic development of the male genital organ presumably by some
genetic event(s). This assumption is not unreasonable, because O. arrowianus murakii inhabit¬
ing the mid-eastern part of the Kinki District has been shown to be a derivative of the hybrid
between O. arrowianus (c?) and O. iwawakianus (-?-) (Imura et al ., 2005b). Note that the
Kongo Mountains are situated just about 60 km west of the habitat of <9. arrowianus murakii ,
suggesting that O. arrowianus once inhabited the intermediate region between the mid-eastern
part of the Kinki District and the Kongo Mountains. The separation of the ND5 gene sequence
between the common ancestor of O. arrowianus in the Chubu District and that of the ancestor of
O. uenoi was calculated to have occurred about 5.6 million years ago. Ohomopterus uenoi is
clearly separable either from O. arrowianus or O. iwawakianus , and is one of the rare cases for
the differentiation of a distinct morphological species from a hybrid-derived ancestor.

Acknowledgements

We express our appreciation to Hideko Kanda for skillful technical assistance. Thanks are
also due to all our friends who have collaborated with us by supplying invaluable specimens and
discussion. This work was supported in part by Grant-in-Aid for Scientific Research (B) (no.
13575013) from Japan Society of the Promotion of Science.

32

Osamu Tominaga et al.

Fig. 7. Genital organ of Ohomopterus kiiensis (d\ Mt. Nagao-yama, Kumano-shi, Mie Pref.; Miyama-
mura, Wakayama Pref.). Captions for each illustration correspond to those in fig. 3. Scale: 2 mm for a,
d-f; 1 mm for b, c, g-i; 0.66 mm for j.

ill • #### • • rn 9S • /Ml * • • -kW£= : 5 i3>K'J7

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References

Hiura, I., 1965. Distribution of Apotomopterus beetles in the Kongo-Ikoma Mountain Range, Central
Kinki, Japan, with special reference to their geohitoric backgrounds (Coleoptera: Carabidae). The
Bulletin of the Osaka Museum of Natural History , (18): 49-68. (In Japanese, with English title and
summary.)

Imura, Y., K. Akita, M. Okamoto, O. Tominaga, N. Kashiwai, Z.-H. Su, T. Ojika and S. Osawa,
2005a. On Ohomopterus arrowianus kirimurai (Coleoptera, Crabidae) as examined by phylogenetic
trees of mitochondrial ND5 gene and nuclear ITS I DNA as well as morphology of genital organs.
The Entomological Review of Japan, 60: 35-38.

Origin of Ohomopterus uenoi

33

Imura, Y., 0. Tominaga, N. Kashiwai, M. Okamoto, Z.-H. Su, T. Ojika, K. Akita and S. Osawa,
2005b. Phylogenetic properties of Ohomopterus iwawakianus (Coleoptera, Carabidae) as evidenced
by the sequence comparisons of mitochondrial ND5 gene and nuclear internal transcribed spacer I:
Extensive participation of O. iwawakianus in the faunal establishment of the genus Ohomopterus in
the Kinki District. Elytra, Tokyo, 33 (in press).

Ishikawa, R., 1960. A new species of Apotomopterus from Japan (Coleoptera, Carabidae). Kontyu, 24:
168-169.

-, 1991. The evolution of Carabus : Divergence and isolating mechanisms. 295 pp. Yasaka Shobo,

Tokyo. (In Japanese.)

Komiya, J., 1971. Classfication of the genus Apotomopterus. Insect Magazine, (76): 22-64. (In Japanese.)

Kinki Research Group of Carabid Beetles (Tominaga, O., K. Katsura, K. Harusawa, I. Hiura, K. Tani
and N. Doi), 1979. Carabid beetles of the Kinki District in the collection of the Osaka Museum of
Natural History. Special Publications from the Osaka Museum of Natural History, 11, 83 pp. (In
Japanese, with English title.)

Okamoto, M., O. Tominaga, Z.-H. Su., Y. Imura, N. Kashiwai, N. Ojika, K. Akita and S. Osawa, 2005.
Differentiation of Ohomopterus yaconinus and its “subspecies” (Coleoptera: Carabidae) inferred from
DNA sequences of mitochondrial ND gene and internal transcribed spacer I (ITS I). Elytra, Tokyo, 33
(in press).

Osawa, S., Z.-H. Su and Y. Imura, 2004. Molecular phylogenyand evolutin of carabid ground beetles.
191 pp. 119 figs. Springer Verlag, Tokyo.

Saito, S., Z.-H. Su, O. Tominaga, N. Kashiwai and S. Osawa, 2003. Pattern of colonization and differ¬
entiation in Ohomopterus albrechti and its related species (Carabinae: Carabidae). The Entomological
Review of Japan, 58: 83-94.

Su, Z-H., O. Tominaga, T. Ohama, E. Kajiwara, R. Ishikawa, T.S. Okada, K. Nakamura and S.
Osawa, 1996. Parallel evolution in radiation of Ohomopterus ground beetles inferred from mitochon¬
drial ND5 gene sequences. Journal of Molecular Evolution, 43: 662-671.

- ,- , M. Okamoto and S. Osawa, 1998. Origin and diversification of hind-wingless

Damaster ground beetles within the Japanese Islands as deduced from mitochondrial ND5 gene
sequences (Coleoptera, Carabidae). Molecular Biology and Evolution, 15: 1025-1039.

-, Y. Imura and S. Osawa, 2001. Evolutionary discontinuity of the carabid ground beetles.

Journal of Molecular Evolution, 53: 517-529.

-,-, M. Okamoto, C.-G. Kim, H.-Z. Zhou, J.-C. Paik and S. Osawa, 2004. Phylogeny and

evolution of Digitulati ground beetles (Coleoptera, Carabidae) inferred from mitochondrial ND5 gene
sequences. Molecular Phylogenetics and Evolution, 30: 152-166.

(Received February 1, 2005; Accepted February 26, 2005)

Em. Rev. Japan , 60 (1): 35-38, April 30, 2005

On Ohomopterus arrowianus kirimurai (Coleoptera: Carabidae) as
Examined by Phylogenetic Trees of Mitochondrial ND5 Gene and
Nuclear ITS IDNA as well as Morphology of Genital Organs

Yuki Imura", Katsumi Akita 2 ', Munehiro Okamoto 3 ’, Osamu Tominaga 4 ',
Nobuo Kashiwai 5 ’, Zhi-Hui Su 6 *', Tooru Ojika 7 ’ and Syozo Osawa 8 '

1) Shinohara-cho, 1249-8, Kohoku-ku, Yokohama, 222-0026 Japan

2) Iba-cho 66, D-304, Hisai City, Mie, 514-1108 Japan

3) Department of Laboratory Animal Science, School of Veterinary Medicine,
Faculty of Agriculture, Tottori University, Tottori, 680-8553 Japan

4) A312, 4-1-15, Shibatsuji-cho, Nara, 630-8114 Japan

5) Hosen-Gakuen High School, Chuo, Nakano-ku, Tokyo, 164-8628 Japan

6) JT Biohistory Research Hall, 1-1 Murasaki-cho, Takatsuki, Osaka, 569-1125 Japan

7) Imahon-machi, 2-9-27-301, Anjo-shi, Aichi, 446-0008 Japan

8) Ushita-Asahi 2-4-7-1003, Higashi-ku, Hiroshima, 732-0067 Japan

Abstract Ohomopterus arrowianus kirimurai , recently described from a very restricted area
of the southeastern periphery of the Kii Peninsula, was investigated by the phylogenetic trees
constructed using the mitochondrial ND5 gene and the nuclear ITS I DNA sequence as well as
morphology of the genital organ of both sexes. The results indicate that this taxon carries the
ND5 gene and the ITS I inseparable from those of the nominotypical O. arrowianus.
However, its female genital organ is of O. iwawakianus- type, which is clearly distinguishable
from that of O. arrowianus- type.

Introduction

Recently, a new subspecies of Ohomopterus arrowianus was described by Kubota and
Yahiro (2003) under the name kirimurai from the southeastern periphery of the Kii Peninsula
based on slight differences in the external and male genitalic morphologies as compared with all
the known races of O. arrowianus. The habitat of this taxon is restricted to an extremely small
area in Mihama-cho near the southern end of Mie Prefecture. In this paper, we wish to present
the results of phylogenetic analyses using the mitochondrial ND5 gene and the nuclear ITS I
DNA sequence, together with the detailed morphologies of the genital organ of both sexes.

*Corresponding author: E-mail: su.zhihui@brh.co.jp

36

Yuki Imura et al.

Results and Discussion

The ND5 gene of O. a. kirimurai in the phylogenetic tree is surely in the cluster I defined
in the previous paper (Fig. 1 of Tominaga et al. 2005), to which the ND5 gene of all the O.
arrowianus specimens belong. This suggests strongly that the ND5 gene of kirimurai was
derived from O. arrowianus , which was immigrated from the southern area of the Chubu
District (presumably the Atsumi Peninsula) at the time when the Ise-wan Bay was still closed.

The ITS I of kirimurai is in the cluster B (Fig. 2a of Tominaga et al ., 2005) together with
that of the nominotypical arrowianus , and not in the cluster A, to which ITS I of all O.
iwawakianus belong.

As shown in Fig. 1, the basic structure of the male genital organ of “subsp.” kirimurai is
almost the same as that of the nominotypical arrowianus , though the figures of the aedeagus and
endophallus were not shown in the original description and are illustrated for the first time in the
present paper. In the nominotypical arrowianus from the Chubu District, the inner plate of the
vaginal apophysis is rather stable in the shape, which is characterized by a chestnut-like or a
reverse heart shaped outline in the dorsal view (see Tominaga et al ., 2005, Fig. 4j), while that
of kirimurai is of O. iwawakianus type (see Fig. lj~m of this paper and Fig. 6j of Tominaga et
al ., 2005), more strictly, of the intermediate between the A-type and B-type of the same species
(Kamiyoshi, 1963; Komiya, 1971). Although O. iwawakianus shows considerable geographical
and individual variations in the shape of this plate, the plate of kirimurai surely resembles that
of O. iwawakianus , and is clearly different not only from O. arrowianus but also from O. kiien-
sis or O. yaconinus, suggesting that kirimurai is not a mere local race of O. arrowianus.

The results of the ND5 gene and ITS I look like as if kirimurai was derived directly from a
pure line of O. arrowianus , despite that kirimurai has the iwawakianus -type female genital
organ. There are at least three possibilities to account for this superficial discrepancy between
the morphology and the molecular data. The first possibility would be that after immigration of
O. arrowianus from the Chubu District to the Mihama-cho of the Kii Peninsula, some muta¬
tion^) leading to the female genital organ of the iwawakianus- type. The second possibility
would be that the ITS I and the gene(s) for the female genital organ are separately localized in
the different chromosomes. Upon the hybridization between O. arrowianus (-?-) and O.
iwawakianus (d*), there should be an opportunity to have conveyed the gene(s) for the O.
iwawakianus female genital organ to the offspring together with the O. arrowianus mitochon¬
dria, having resulted in establishment of kirimurai , although this event is unable to detect by the
ITS I analysis alone. The third possibility would be that the ITS I and the gene(s) for the female
genital organ coexisted on the same chromosome, and upon the hybridization of O. arrowianus
(■¥■) with O. iwawakianus (<?), a recombination between O. arrowianus and O. iwawakianus
occurred so as to have produced a chimera recombinant carrying the ITS I of O. arrowianus and
the gene(s) of O. iwawakianus female genital organ. If such an event took place, kirimurai hav¬
ing the ND5 gene and ITS I of the arrowianus-type would have resulted. It is necessary to iden¬
tify and localize the ITS I and the gene(s) responsible for the morphology of female genital
organ on the chromosome(s).

In summary, O. arrowianus kirimurai is a fortuitous remnant with an accompanied muta¬
tional change(s) for the female genital organ, or a descendant resulted by chance from a hybrid
between O. arrowianus arrowianus (-?-) and O. iwawakianus (cT). At present, there is no evi-

Origin of Ohomopterus arrowianus kirimurai

37

Fig. 1. Genital organ of Ohomopterus arrowianus kirimurai (Mihama-cho, Mie Pref.). a, Aedeagus in right
lateral view; b, apical part of aedeagus in the same view; c, ditto in dorsal view; d, fully everted
endophallus in right lateral view; e, ditto in posterior view; f, ditto in anterior view; g, copulatory piece in
ventral view; h, ditto in lateral view; i, peripheral rim of gonopore in right lateral view; j-m, inner plate
of vaginal apophysis in dorsal view. Scale: 2 mm for a, d-f; 1 mm for b, c, g-i, 0.7 mm for j-m.

dence to decide which possibility is more likely. The quite narrow distributional range of this
“subspecies” does not warrant optimism for its long survival.

Acknowledgements

We express our appreciation to Hideko Kanda for skillful technical assistance. Thanks are
also due to Mr. Naoaki Toda (Nagoya) and all our friends who have collaborated with us by
supplying invaluable specimens and discussion. This work was supported in part by Grant-in-
Aid for Scientific Research (B) (no. 13575013) from Japan Society of the Promotion of Science.

38

Yuki Imura etal.

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References

Kamiyoshi, M., 1963. On the female genitalia of Apotomopterus from Japan (Col., Carabidae). Insect
Science, 13: 1-8. (In Japanese.)

Komiya, Z., 1971. Classification of the genus Apotomopterus. Insect Magazine, (76): 22-64. (In
Japanese.)

Kubota, K. and K. Yahiro, 2003. Description of an isolated and specialized population of Carabus
arrowianus (Breuining, 1934) (Coleoptera, Carabidae) discovered in the southernmost part of Mie
Prefecture, Japan, as a new subspecies, with analyses of its morphological features. Biogeography, 5:
9-15.

Tominaga, O., Y. Imura, M. Okamoto, Z.-H. Su, T. Ojika, N. Kashjwai. and S. Osawa, 2005. Origin of
Ohomopterus uenoi (Coleoptera: Carabinae: Carabidae) as deduced from comparisons of DNA
sequences of mitochondrial ND5 gene and nuclear internal transcribed spacer I (ITS I) with morpho¬
logical characters. The Entomological Review of Japan, 60: 23-33.

(Received February 1, 2005; Accepted February 21, 2005)

Ent. Rev. Japan , 60 (1): 39-51, April 30, 2005

Five New Species of the leptopus Group of
Harpaline Genus Trichotichnus (Coleoptera: Carabidae)
from Central and Northeastern Japan,
with Note on Taxonomic Position of T. tsurugiyamanus

Noboru Ito

1-7-18 Higashiuneno, Kawanishi City, Hyogo Pref., 666-0117 Japan

Abstract Five new species of the leptopus group of the genus Trichotichnus are described
under the names of Trichotichnus marubayashiorum from Kamikochi and the adjacent places,
T. iidesanus from Mt. Iide-san and near regions, T. choukaisanus from Mt. Choukai-san, and
T. mizunoi and T. hosodai from Gozaishi Spa. Further, it is indicated by detailed reexamina¬
tion that T. tsurugiyamanus Habu belongs to the congruus group.

In 1973, the leptopus group of the genus Trichotichnus Morawitz was established by Habu
and known species were reviewed with description of a new species. After that, several new
species were described by Habu and Kasahara. In 1996,1 revised all the species from western
Japan including Fukui Pref., with descriptions of some new species. In next year, Morita treat¬
ed all species of the group including many new species described then and proposed six com¬
plexes. The species of the group are well diversified locally owing to vestigial hind wings, and
many unknown species may be estimated to occur yet. I collected in Central and Northeast
Japan and found additional new species.

Habu (1973) included Trichotichnus tsurugiyamanus Habu within the leptopus group in his
revision and I followed his treatment (Ito, 1996). Recently I reexamined characteristics of
pronotum, legs, 6th abdominal sternite and male genitalia of tsurugiyamanus and concluded that
tsurugiyamanus should be transfer to the congruus group.

In this paper I am going to describe five new species of the leptopus group as following:
Trichotichnus ( Trichotichnus) marubayashiorum from Kamikochi and adjacent places, T. (T.)
iidesanus from Mt. Iide-san and nearby regions, T. ( T .) choukaisanus from Mt. Choukai-san, T.
(T.) mizunoi and T. (T.) hosodai from Gozaishi Spa at the foot of Mt. Houou. Also T. (T.) tsu¬
rugiyamanus is newly transferred to the congruus group.

Before going further, I would like to express my hearty thanks to Mr. Yoshiyuki Ito, Kochi,
Mr. Kozo Mizuno, Uji, Mr. Kouichi Hosoda, Nirasaki, Mr. Masato Mori, Nishinomiya, Drs.
Shin-ichirou Yoshimatsu and Kazuhiko Konishi of the National Institute of Agro-Environ-
mental Sciences, Tsukuba, and Dr. Shuhei Nomura of the National Science Museum (Nat.
Hist.), Tokyo for their kindly offering or loaning material. Also, I heartily thank Mr. Seiji
Morita, Tokyo, for his valuable opinion. Lastly, I am indebted to parents’ family of my wife,
Marubayashi for their kind support on my field work in Nagano Pref.

40

Noboru Ito

I employ the abbreviation of depository as following: the National Institute of Agro-
Environmental Sciences as NIAS; the Osaka Museum of Natural History as OMNH; the
National Science Museum (Nat. Hist.) as NSMT; the author’s collection as NIc. Concerning
measurement of body parts, see my former paper.

Figs. 1-5. Habitus of Trichotichnus spp.-1, Trichotichnus ( Trichoticlmus ) marubayashiorum N. Ito, sp.

nov.; 2, T. ( T .) iidesanus N. Ito, sp. nov.; 3, T. ( T .) choukaisanus N. Ito, sp. nov.; 4, T. ( T .) mizunoi N.
Ito, sp. nov.; 5, T. ( T .) hosodai N. Ito, sp. nov.

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

41

Fig. 6. Male genitalia of Trichotichnus (Trichotichnus ) marubayashiorum N. Ito, sp. nov.
d, dorsal aspect; v, ventral aspect. Scale: 1 mm.

Trichotichnus ( Trichotichnus ) marubayashiorum N. Ito, sp. nov.

(Figs. 1, 6)

Body rather wide, oblong, black, shiny, with iridescent lustre on elytra; labial and maxil¬
lary palpi, antennae and legs somewhat light to dark reddish brown, labrum and mandibles
brownish black.

Head gently convex, a little large, 0.65-0.69 times as wide as pronotum, with several
punctures on frons; labrum triangularly concave at apex; clypeus ante-trapezoidally emarginate
at apex, depressed in apical part, weakly elevated behind there; clypeal suture shallow but clear;
frontal impressions moderately deep throughout, reaching supraorbital grooves; interocular
space wide, 0.71 times as wide as width of head including eyes; eyes not convex, short; temples
long, 0.4 times of eye length, straightly convergent behind; genuine ventral margins of eyes nar¬
rowly separated from buccal fissure; labial palpi slender, 2nd segment slightly longer than the
3rd; ligula wide, acutely produced laterad at apical angles; paraglossae narrow, slightly pro¬
longed behind ligula; mentum with epilobes strongly widened apicad, median tooth rounded at

42

Noboru Ito

apex; microsculpture obscure, consisting of isodiametric meshes near clypeal apex, and partly
of transverse meshes on frons.

Pronotum 1.33-1.39 times as wide as long, fairly convex, wholly covered with punctures,
which are fine on central area and become coarser towards surrounding areas, especially coarse
in basal foveae; apex uniformly emarginate, with border obscure in middle; sides clearly arcuate
apicad and gently oblique basad from the widest point, rather deeply sinuate before base; base
one-tenth wider than the apex, barely bisinuate, and thickly bordered lengthwise; apical angles
narrowly rounded; basal angles rectangular, produced laterad at tips; lateral furrows narrow in
apical third, thence gradually and weakly widened backwards, fallen into basal foveae which are
large and longitudinally grooved at inner side; front transverse impression rather deep, the hind
one indistinct; median line thin but clear; microsculpture clearer in female than in male, consist¬
ing of mixture with isodiametric and transverse meshes.

Elytra widely suboval, widest at apical two-fifths, 1.40-1.50 as long as wide, one-fourth to
three-tenths wider than pronotal width, gently and uniformly convex, without punctures; bases
shallowly emarginate (almost straight in one specimen), humeral angles weakly produced for¬
wards, with a tiny tooth at each tip; sides feebly concave in basal fifth, very shallow in preapical
sinus; apices narrowly rounded at tips; striae narrow and shallow, scutellar striole short; inter¬
vals weakly convex, becoming a little more convex basad and apicad, a setiferous pore of 3rd
interval situated slightly before middle; marginal series composed of 23-28 umbilicate pores,
continuous though rather wide at intervals between several neighboring pores in middle;
microsculptures invisible or partly visible as vague transverse lines. Hind wings well reduced,
one-fifth of elytral length.

Metepistema 0.81 times as long as wide; 6th abdominal sternite truncate or feebly notched
in male and widely rounded in female at apex, unisetose in male and bisetose in female at each
side.

Legs long; hind femora bisetose along hind margin; fore tibiae sulcate along full length,
trispinose apico-external margin; hind tarsi 1.13 times in male and 1.05 times in female as long
as the width of head, relative length of 1st to 4th segments = 1.00 : 0.75 : 0.58 : 0.38, claw seg¬
ment trisetose along inner margin and tri- or quadrisetose along the outer one.

Aedeagus (Fig. 6) thick, gradually thinned apicad, slightly curved ventrad at tip; apical
orifice wide, directed oblique-dorsad, armed with a copulatory piece peg-shaped; apical orifice
elongate-triangular, thinly bordered; ventral surface gently curved, weakly arcuate in middle.

Length: 10.9-11.9 mm. Width: 4.5-4.9 mm.

Holotype: <?, Kamikochi, Nagano, 27—30. VI. 1958, H. Yokoyama leg. Paratypes: 1
same locality as the holotype, 27. VI. 1958, H. Yokoyama leg.; 2££, same locality as the
holotype, 21. VII. 1959, T. Shibata leg.; 1 £, Iwanadome, Nagano, 5. VIII. 1965, Y. Imai leg.;
1 c?, Tokugo Pass, Nagano, 11. VIII. 1992, N. Ito leg. (preserved in NIc); 3<?<T, Hirayu pass,
Gifu, 20. VII. 1988, M. Saito leg. (preserved in NSMT and NIc).

Remark: This new species is allied to Trichotichnus ( Trichotichnus ) furihatai Morita
from Nakabusa-onsen, but the body is robuster, eyes are flatter, and the aedeagus is thinner at
apical lobe in lateral aspect.

Etymology: The new species, “ marubayashiorum ” is named after Marubayashi family.

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

43

Fig. 7. Male genitalia of Trichotichnus ( Trichotichnus) iidesanus N. Ito, sp. nov.
d, dorsal aspect; v, ventral aspect. Scale: 1 mm.

Trichotichnus ( Trichotichnus ) iidesanus N. Ito, sp. nov.

(Figs. 2, 7)

Body moderate-sized, gently convex, black, shiny, weakly iridescent on elytra; palpi,
antennae and legs yellowish brown to a little dark brown, and labrum and middle of mandibles a
little darker.

Head more or less large, 0.68-0.73 times the pronotal width, fairly raised on vertex, some¬
what coarsely and sparsely punctate, with wide interocular space three-fourths of the width of
head; labrum shallowly notched at apex; clypeal suture thin, uninterrupted; frontal impressions
almost straightly divergent behind, reaching supraorbital grooves, rather deep throughout; eyes
slightly prominent; temples long, one-third of the eye length, rather well tumid; space between
genuine ventral margins of eyes and buccal fissure; antennae slender, reaching basal sixth of
elytra, 3rd segment as long as the 4th and twice the 2nd ; labial palpi short; ligula and paraglos-
sae similar in shape to the previous new species; epilobes of mentum weakly widened; micro¬
sculpture a little clearer in female than in male, composed of isodiametric meshes on apical area
of clypeus and partly of transverse meshes on frons.

44

Noboru Ito

Pronotum somewhat transversely cordate, widest at apical third, 1.40-1.46 times as wide
as long, gently declivous apico-laterad; sides a little weakly arcuate from apex to middle, thence
straightly oblique backwards, sinuate before base; apex rather strongly emarginate, with border
vague in middle; base one-tenth wider than apex, shallowly emarginate, feebly rounded at the
sides, and thickly bordered throughout; apical angles fairly produced forwards, narrowly round¬
ed; basal angles acute, triangularly protruding laterad; lateral furrows narrow, weakly expanded
backwards; basal foveae moderate in size, weakly humped in the middle, longitudinally grooved
inside the humps; front transverse impressions rather deep, the hind one obscure; median line
fine, though somewhat clear, lying between the impressions; surface very sparsely and finely
punctate on central area, a little more coarsely and moderately so in apical and lateral areas,
rather coarsely so in lateral furrows and basal foveae where the punctures are dense and partly
confluent; microsculpture a little more clearly observed in female than in male, partly visible as
mixtures with isodiametric and transverse meshes.

Elytra oval, rather short, 1.39-1.45 times as long as wide, gently raised, very sparsely with
very minute punctures; sides not concave, uniformly arcuate or straight behind humeral arc,
somewhat deeply sinuate before apices which are rather produced and narrowly rounded at dis¬
tal margins; bases shallowly emarginate, acute at shoulder angles; striae thin, shallow, and fine¬
ly crenulate; scutellar striole short; intervals flat on disc and becoming a little convex apicad,
basad and laterad, a setiferous dorsal pore of 3rd interval situated between apical four- and five-
ninths; marginal series continuous, consisting of 24-30 umbilicate pores. Hind wings obliterat¬
ed.

Metepisterna 0.90 times as long as wide; 6th abdominal sternite in male weakly arcuate at
apical margin and unisetose at each side and in female a little more strongly arcuate and bise-
tose.

Legs rather long; hind femora bisetose along hind margin; fore tibiae sulcate lengthwise,
apico-extemally with three spines; hind tarsi in male 1.02-1.08 times and in female 0.95-0.96
times as long as the width of head, relative length of 1st to 4th segments as 1.00 : 0.70 : 0.50 :
0.31, ventral sides of claw segment trisetose along external margin and tri- or quadrisetose
along inner one.

Aedeagus (Fig. 7) stout, abruptly tapered forwards, thinned at apex, feebly curved
obliquo-ventrad at tip; apical orifice almost directed dorsad, widely open, armed with a short
copulatory piece peg-shaped; apical orifice short, widely rounded and bordered at distal margin;
ventral surface not curved.

Length: 9.9-11.0 mm. Width: 4.2-4.7 mm.

Holotype: <?, Nukumidaira, Mt. Iide, Yamagata, 14. VIII. 1984, N. Ito leg. (preserved in
OMNH). Paratypes: 3<?V, 2£ Y, same data as the holotype (preserved in NIc); 2
Kurokawa, Niigata, 24. VII. 1957, K. Baba leg.; 1 <3\ ditto, 27. VII. 1957; 1-?-, ditto, 17. IX.
1957; 1 <?, Kanamaru, Niigata, 2. VIII. 1957, K. Baba leg. (preserved in NIAS and NIc).

Remark: This new species is similar in lateral aspect of aedeagus to Trichotichnus
armiger Morita, but being judged from the original description of the latter, the new species is
distinguished from it by the eyes flatter, the elytra wider, and the aedeagus bearing apical lobe
shorter and bordered at distal margin.

Etymology: The species “ iidesanus ” is named after the type locality, Mt. Iide-san.

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

45

Fig. 8. Male genitalia of Trichotichnus ( Trichotichnus ) choukaisanus N. Ito, sp. nov.
d, dorsal aspect; v, ventral aspect. Scale: 1 mm.

Trichotichnus ( Trichotichnus ) choukaisanus N. Ito, sp. nov.

(Figs. 3, 8)

This new species is allied to Trichotichnus ( Trichotichnus ) abei Morita, but the body is a
little narrower, the pronotum is thinner in lateral margins and narrower in lateral furrows, and
the aedeagus bears apical orifice wider and truncate at distal margin.

Body slightly brownish black, iridescent on elytra; head brownish, maxillary and labial
palpi, antennae, and legs light brown. Head wide, 0.70—0.72 times as wide as pronotal width,
fairly convex, with very sparse and somewhat coarse punctures; eyes relatively large, rather
convex; clypeal suture more or less deep; frontal impressions relatively deep, shallowed behind;
3rd segments of antennae slightly longer than the 4th and twice the 2nd; ligula and paraglossae
alike former new species, T. iidesanus ; 2nd segment of labial palpi one-ninth longer than the
3rd; median tooth rounded; microsculpture obscure. Pronotum widest at apical third, 1.40-1.44
times as wide as long, rather convex; sides weakly arcuate from apex to basal third, thence lin¬
early oblique and sinuate before base; apex gently and a little strongly emarginate; base one-
sixth wider than apex, weakly bisinuate-emarginate, thickly bordered; surface smooth on central
area, sparsely and minutely punctate in apical area, coarsely and densely in lateral furrows and
basal foveae; basal angles rectangular or feebly protruding laterad; microsculpture vague, partly
observed as transverse meshes in basal foveae. Elytra oval, wide, two-fifths longer than wide.

46

Noboru Ito

one-third wider than pronotal width, weakly and uniformly convex, wholly smooth; a setiferous
pore on 3rd interval situated between basal two-hfths and three-sevenths; marginal series inter¬
rupted medially, composed of (10-11) + (12-13) umbilicate pores. Hind wings vestigial, one-
sixth of elytra! length. Metepisternuna 0.93 times as long as wide; 6th abdominal segment in
male truncate and in female rounded at apex. Legs not long; hind tarsi in male as long as and
one-tenth in female shorter than the width of head, relative length of 1st to 4th segments as fol¬
lowing: 1.00 : 0.70 : 0.50 : 0.32, chaetotaxy of claw segment same as that of former new
species, T. iidesanus. Aedeagus (Fig. 8) moderately stout, gradually thinned apicad, thickened at
tip; apical orifice wide and sinuately truncate at apical margin, copulatory piece very small and
conical; ventral surface straight, slightly curved ventrad.

Length: 9.3-9.9 mm. Width: 4.CM-.2 mm.

Holotype: d\ Mt. Choukai, Yamagata, 12. VIII. 1985, N. Ito leg. (preserved in OMNH).
Paratypes: 1 Y, same data as the holotype; 1 same locality as the holotype, 21-22. VII. 1957,
T. Hori leg. (preserved in NIc).

Etymology: This specific name is derived from the type locality, Mt. Choukai-san.

Trichotichnus (Trichotichnus) mizunoi N. Ito, sp. nov.

(Figs. 4, 9)

Body oblong, rather elongate, black, shiny, with iridescent lustre on elytra; maxillary and
labial palpi, antennae and legs reddish brown, middle of mandibles and frons slightly brownish.

Head large, 0.70 times as wide as pronotal width, fairly elevated on frons, with vague and
very sparsely punctures; labrum deeply and triangularly emarginate at apex; clypeal suture and
frontal impressions alike three former new species; eyes flat or weakly prominent; temples
somewhat developed, one-third of eye length; antennae missing apical five segments, relative
length of 2nd, 3rd and 4th segments as 1.00 : 2.00 : 1.90; mentum with median tooth small and
shaip, epilobes not widened forwards; microsculpture relatively clear, detected as isodiametric
meshes in apical area of clypeus and as mixtures with fine square and isodiametric meshes on
frons and vertex.

Pronotum cordate, fairly reflected baso-laterally, weakly convex, widest at apical third,
1.44 times as wide as long; sinus before base shallow or rather deep; apex well emarginate,
unbordered in middle; base not wide, one-tenth wider than apex, almost straight, feebly arcuate
at sides, thickly bordered throughout; apical angles very narrowly rounded; basal angles acute,
not or weakly produced laterad at tips; lateral furrows somewhat wide, abruptly expanded basad
from apical two-fifths; basal foveae almost flattened, grooved at inner side; both front and hind
transverse impressions relatively deep; median line fine and clear, obliterated near both the
impression; microsculpture finely and clearly impressed as transverse meshes.

Elytra rather narrowly oval, nearly a half longer than wide, 1.23 times as wide as pronotal
width, weakly convex, without punctures; sides smoothly and gently arcuate behind humeri,
weakly sinuate before apices which are very narrowly rounded at tips; bases slightly emar¬
ginate, angularly meeting with lateral margins; striae relatively deep and finely crenulate, scutel-
lar striole moderately short; intervals weakly raised even on disc, a setiferous pore on 3rd inter¬
val situated between middle and apical three-fifths; marginal series consisting of 24-27 umbili¬
cate pores; surface obscurely and finely microlined. Hind wings a little more weakly reduced

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

47

Fig. 9. Male genitalia of Trichotichnus ( Trichotichnus ) mizunoi N. Ito, sp. nov.
d, dorsal aspect; v, ventral aspect. Scale: 1 mm.

than usual, one-fourth of the elytral length.

Metepisterna 0.93 times as long as wide. Legs long; hind femora bisetose; fore tibiae sili¬
cate, trispinous along apico-external margin; hind tarsi long, 1.20 times in male and 1.10 times
in female as long as the width of head, relative length of 1st to 4th segments as 1.00 : 0.70 : 0.55 :
0.42, claw segment ventrally tri- or quadrisetose along inner margin and trispinose along the
outer one.

Aedeagus (Fig. 9) stout, gradually tapered forwards from middle, thin at apex, slightly
reflected at tip, with large basal bulb; apical orifice widely open, weakly sinuate at distal mar¬
gin, armed with a copulatory piece peg-shaped, long and robust; apical lobe subtrapezoidal, bor¬
dered and weakly rounded at distal margin.

Length: 12.0-12.5 mm. Width: 4.9-5.1 mm.

Holotype: <?, Gozaishi Spa, Yamanashi, 6. VII. 1992, H. Hosoda leg. (preserved in
OMNH). Paratype: 1 same data as the holotype (preserved in NIc).

Remark: This new species is peculiar in having long and robust copulatry piece on inner
sac of aedeagus. The species is similar in shape of body to Trichotichnus ( Trichotichnus ) hiran-

48

Noboru Ito

Fig. 10. Male genitalia of Trichotichnus ( Trichotichnus ) hosodai N. Ito, sp. nov.
d, dorsal aspect; v, ventral aspect. Scale: 1 mm.

ishii Morita from Hakone, Tanzawa and Mt. Fuji, and it is difficult to distinguish both species
by external characteristics, but in new species the pronotum is more weakly sinuate before base
and more narrowly covered with punctures, and hind tarsi are shorter.

Etymology: The specific name “ mizunoi ” is dedicated to Mr. Kozo Mizuno who offers
many invaluable material for my study.

Trichotichnus ( Trichotichnus) hosodai N. Ito, sp. nov.

(Figs. 5, 10)

This new species is very similar to Trichotichnus ( Trichotichnus ) daibosatsunis Kasa-
hara, but the pronotum is a little narrower in lateral furrows, weakly humped in basal foveae
instead of being flattened there and more deeply sinuate before base, the tarsi are wider, and the
aedeagus bears copulatry piece wider and acute at its tip instead of rounded.

Body oblong, relatively narrow, black, shiny, iridescent on elytra; maxillary palpi, anten¬
nae, and legs light brown, mandibles and head slightly brownish. Head large, 0.71-0.76 times as
wide as pronotal width, with narrow interocular space two-thirds of the width of head; eyes
comparatively larger and rather prominent than usual species of the leptopus group; antennae

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

49

Fig. 11. Male genitalia of Trichotichnus ( Trichotichnus) tsurugiyamanus Habu.
d, dorsal aspect. Scale: 1 mm.

slender, long, reaching basal fifth of elytra, epilobes of mentum weakly widened apicad. Pro-
notum clearly cordate, rather well convex, widest at apical third, one-third wider than long;
sides not so strongly sinuate before base; base almost truncate, narrow, 1.14-1.16 times as wide
as apex; basal angles rectangular; microsculpture vaguely and partly visible as transverse mesh¬
es. Elytra rather well convex, suboval to oblong, approximately a half longer than wide, widest
between apical five-ninths and half; sides barely concave behind humeri; apices somewhat
widely rounded at distal margins; marginal series continuous, composed of 25-28 umbilicate
pores. Hind wings relatively long, three-fourths of the elytral length. Legs long; fore tibiae
rather clearly sulcate; hind tarsi 1.11-1.16 times in male and 1.03 times in female as long as the
width of head. Aedeagus (Fig. 10) not robust, almost straightly prolonged, thickened at apex;
inner sac not wide, armed with copulatory piece widely conical and pointed at its tip; apical lobe
subquadrate, longer than wide.

Length: 10.0-11.0 mm. Width: 4.1-4.4 mm.

Holotype: cT, Gozaishi Spa, Yamanashi, 20. VII. 1990, K. Hosoda leg. (preserved in
OMNH). Paratypes: 1 <?, l£, same locality and collector of the holotype, 28. VI. 1990 (pre¬
served in NIc).

Etymology: This specific name is dedicated to Mr. Kouichi Hosoda, Nirasaki, who is an
excellent collector of insects and captured specimens of the type series.

50

Noboru Ito

Trichotichnus ( Trichotichnus ) tsurugiyamanus Habu

(Fig. 11)

Trichotichnus tsuriigiyamcinus Habu. 1957, Kontyu, 27: 131. 1961, Bull. Natn. Inst. Agric. (C), 13: 154.

1973, Fauna Japnica: 289 (leptopus group).

Trichotichnus tsurugiyamanus. Ito, 1996, Ent. Rev. Japan, 51: 132.

Trichotichnus tsurugiyamanus. Morita, 1997, Elytra, 25: 584.

Specimens examined: 1 c?, Kuwadaira, Minokoshi, Mt. Tsurugi, Tokushima Pref., 3. V. 1958, N.
Kawano leg.; 1 <?, Mt. Yahazu, Awa, 5. VIII. 1961, N. Kawano leg.; 1_, Mt. Tsurugi, Tokushima Pref.,
30. VII. 1960, T. Shibata leg.; 3<? cT, 4-?- ?, Minokoshi, Mt. Tsurugi, Tokushima, 10-11. VI. 2000, N. Ito
leg.; 5^^, 13ditto, 24. VII. 2004; 3<?c?, 1-?-, Mt. Tsurugi, Higashiiyayama-mura, alt. 1,400 m, 8.
VI. 2002, M. Mori leg.; 1 cT, Mt. Takanosu-yama, Hongawa-mura., Tosa-gun, Kochi, 7-9. VIII. 1998, Y.
Ito leg.

Habu (1973) established the leptopus group of the genus Trichotichnus including this
species. I followed this treatment (Ito, 1996). After that, Morita personally suggested me that
the species might not belong to the group, but to the congruus species group. As the result of
my reexamination, it could be made clear as following: 1, pronotum is not cordate, though
transverse, and with punctures more scarce; 2, metepiterna are shortened, though longer than
usual species of the leptopus group; 3, legs are shorter than those of the leptopus group’s
species; 4, foretibiae are never sulcate; 5, sixth abdominal sternite of male is bisetose at each
side; 6, aedeagus (Fig. 11) is straightly tapered distad from basal bulb and with a copulatory
piece sinuate instead of being almost straight or conical. As these characteristics agree to those
of the congruus group. I would like to propose to transfer T. tsurugiyamanus from the leptopus
group to the congruus group.

Distributional pattern of the leptopus group: The species of the group are diversified
very well in narrow areas owing the regression of hind wings. On the other hand, two or more
species often occur sympatrically in quite same region, for examples, Kamikochi, Gozaishi Spa,
the mountainous areas of Kii peninsula, Shikoku district and Kwanto district, Ryuto-zan in
Shizuoka, Daibosatsu in Yamanshi, and Mt. Hiko-san in Fukuoka, etc. Those sympatric species
are genetically isolated to each other and hybrid species are not founded. This phenomenon sug¬
gests that different ancestors invaded to the regions in different ages. Detailed taxonomic study
and distribution data of species from the not surveyed areas are needed further for the analysis
of the phylogenetic relationship of all species.

J: O'JlUfc B Genus Trichotichnus (7 *Y H'>I) CO leptopus group

5 frflio J :Xf T. tsurugiyamanus - Genus Trichotichnus (7*Y

n'^HyS) co leptopus group (i±*^ ? 1973^UilJlx LX i 1996^^^117*15

L^, Trichotichnus

marubayashiorum (v;W^v7t 7 A 7) , III % SI EffiH fr b T. mizunoi ( ^ X 7 7
-Y zf^E ny) £ X T. hosodai (*7*77 *Y^^Ay), \kfrb T. iidesanns

New Species of the Trichotichnus leptopus Group and Note on T. tsurugiyamanus

51

(4 4 ? A '>), ffelll b T. choukaisanus a ^ fj >f y *7 U's)

#¥**egs«>umjM? f %m<nm&\u, iwmsKiij,

ttS:[Ilic*3/'oT5}-^^£ltYj; t §>&£>■£&. ZtztbClt,

-n, MtttwAt, nfflttuftj

tsurugiyamanus (7iVf ? Ay) < congruus group UpjfSi -

li congruus group Ltz.

References

Habu, A., 1957. New species collected by Professor M. ChCijo on Mt. Tsurugiyama in Shikoku, Japan
(Coleoptera, Carabidae). Kontyu, 27: 126-133.

-, 1961. Revisional study of the species of the Trichotichni, the subtribe of the tribe Harpalini,

from Japan (Coleoptera, Carabidae). The Bulletin of the National Institute of Agricultural Sciences
(C), 13 : 127-169.

-, 1973. Carabidae: Harpalini (Insecta: Coleoptera). Fauna Japnica. xii + 430 pp., Keigaku

Publishing, Tokyo.

Ito, N., 1996. Species of the leptopus group of the genus Trichotichnus from western Japan including
Fukui Pref. (Coleoptera: Carabidae: Harpalini). The Entomological Review of Japan, 51 : 121-134.

Kasahara, S., 1991. A new species of the genus Trichotichnus (Coleoptera, Carabidae) from central
Honshu, Japan. Elytra, Tokyo, 19 : 111-114.

Morita, S., 1997. The group of Trichotichnus leptopus (Coleoptera, Carabidae) of Japan. Elytra , Tokyo,
25 : 521-585.

(Received January 29, 2005; Accepted February 22, 2005)

Ent. Rev. Japan , 60 (1): 53-54, April 30, 2005

Replacement Names for the Junior Homonyms of
Two Species of Trichotichnus and a Genus of Harpalini
(Coleoptera: Carabidae: Harpalini)

Noboru Ito

1-7-18 Higashiuneno, Kawanishi City, Hyogo Pref., 666-0117 Japan

Trichotichnus ( Trichotichnus ) pictipennis N. Ito, nom. nov.

Trichotichnus ( Trichotichnus) maculipennis N. Ito, 1998, Bull. Osaka Mus. nat. Hist., 52: 52 (nec
Trichotichnus maculipennis Baehr, 1997).

Trichotichnus maculipennis N. Ito, which was described in 1998, is a homonym of T.
maculipennis Baehr, 1997. So, I would like to give a new name, Trichotichnus pictipennis , to
the former species.

Trichotichnus ( Amaroschesis ) yukii N. Ito, nom. nov.

Trichotichnus ( Amaroschesis) imurai N. Ito, 2002, Ent. Rev. Japan, 57: 182 [nec Trichotichnus
{Trichotichnus) imurai Morita, 1997, Elytra, Tokyo, 25: 539].

Trichotichnus imurai N. Ito, which was described in 2002, is a homonym of T. imurai
Morita, 1997. Therefore I would like to replace a new name, Trichotichnus yukii , to the former.

Genus Merklophonus N. Ito, nom. nov.

Merklia N. Ito, 2004, Jpn. J. Syst. Ent., 10: 107 (nec Merklia Chen, 1997, Ent. Sinica, 4: 306)

Merklia N. Ito described in 2004 is a homonym of lagriid genus Merklia Chen, 1997. I
would like to give a new generic name, Merklophonus , to the former. Dr. John K. Page of the
Thomson Zoological Ltd. informed me that Merklia was already preoccupied by the generic
name of Lagriidae in 1997.1 heartily thank him for his kind support.

References

Baehr, M., 1997. A new species of Trichotichnus Morawitz from northern Australia (Insecta,
Coleoptera, Carabidae, Harpalinae). SPIXIANA, 20: 131-135.

54

Noboru Ito

Chen. B., 1997. Two new genera and two new species of Lagriidae (Coleoptera) from China.
Entomologia Sinica , 4: 306-311.

Ito, N., 1998. Three new species of the genus Trichotichnus from Seram Is., the Moluccas with a note of
distribution (Coleoptera: Carabidae: Harpalini). The Bulletin of the Osaka Museum of Natural
History , 52: 49-56.

-, 2002. Three new species of the subgenus Amaroschesis of the genus Trichotichnus (Coleptera:

Carabidae: Harpalni) from Sichuan, with new record. The Entomological Review of Japan , 57:
181-190.

- , 2004. A new genus and species of the Selenophori group of tribe Harpalini from India

(Coleoptera, Carabidae). The Japanese Journal of Systematic Entomology , 10: 107-112.

Morita, S., 1997. The group of Trichotichnus leptopus (Coleoptera, Carabidae) of Japan. Elytra , Tokyo ,
25: 521-585.

(Received February 22, 2005; Accepted March 4, 2005)

Ent. Rev. Japan , 60 (1): 55-58, April 30, 2005

Propsephus nanshanus, a New Species of Elateridae (Coleoptera)

from Taiwan

Hisayuki Arimoto

Tedukayama-nishi 3^4—21, Sumiyoshi-ku, Osaka, 558-0052 Japan

and

Sergio Riese

Corso Sardegna 46-11 sc. D, 16142, Genova, Italy

Abstract A new species of the elaterid genus Propsephus Hyslop, 1921, is described from
Nantou Hsien, Taiwan, under the name of P. nanshanus.

The genus Propsephus Hyslop, 1921, was established on Psephus beniniensis Candeze,
1859, and includes a mumber of species from the African and the Malagasy Regions and a few
from the Oriental Tropics at present (Schenkling, 1927). We have found a new species of the
genus in the course of our recent study on Elateridae of Taiwan. In this paper, we are going to
describe it as the first species of the genus from Taiwan.

Before going further, we wish to express our sincere gratitude to Dr. Hitoo Ohira, Okazaki,
for his constant guidance and Dr. Hisashi Ashida, Ibaraki, for his critically reading the manu¬
script.

The holotype is preserved in the collection of the Osaka Museum of Natural History.

Propsephus nanshanus sp. nov.

(Figs. 1-4)

Male. Length about 18.5 mm and width 5.5 mm. Body shining, robust, moderately elon¬
gate, almost parallel-sided and convex above; dark brown except for ventral surface, antennae
and legs more or less paler than dorsal surface. Dorsal surface clothed with recumbent, golden-
yellow and rather long setae on head and pronotum but a little shorter on elytra; ventral surface
pale yellow, with recumbent setae.

Head subquadrate, with frons weakly and broadly impressed between eyes, clearly
impressed between antennae; surface coarsely and densely punctate, each puncture seemingly
umbilicate; frontal margin of clypeus rounded and clearly impressed at the middle, but conspic¬
uously ridged above antennal insertions; labrum moderately convex, semicircular and coarsely
punctate; front-clypeal area transverse, broad and somewhat narrowed at the middle. Antennae

56

Hisayuki Arimoto and Sergio Riese

Fig. 1. Habitus of Propsephus nanshanus sp. nov., holotype.

short, not attaining to posterior angle of pronotum, basal segment robust and subclavate; the
second short, subquadrate and about 0.9 times as long as its width; the third obconical and about
2.1 times as long as the second; the fourth elongate-triangular and about 1.4 times as long as the
third; the fourth to tenth normally serrate; apical segment oblong-ovate and about 3.5 times as
long as wide (Fig. 2).

Pronotum trapezoidal, about 1.2 times as long as basal width; sides almost straight in basal
half, gently convergent towards middle, then feebly arcuate and clearly convergent towards
anterior angles; disc gently convex, coarsely and densely punctuate, each puncture umbilicate;
median line smooth and longitudinal, and clearly visible in basal half; posterior angles project¬
ing postero-laterad, each with a distinct carina above. Scutellum lingulate and obtusely pointed;
surface coarsely punctate and clearly convex in apical third.

Elytra about 2.3 times as long as its basal width, with sides almost parallel in basal three-
fifths, then rounded and gradually convergent towards apices; striae definite, bearing elongate
punctures; intervals feebly elevated, sparsely and unevenly punctate and transversely rugose.

Legs stout; apical end of the second tarsal segments distinctly membranous-lobate ventral-
ly; the third more noticeably lobate than the second; claws simple.

Propleura rather shallow, densely and umbilicate-punctate in each apical three-fourths, but
the punctures are slightly smaller than those of pronotum. Prosternum densely and umbilicate-
punctate, each punctures clearly larger than in propleura. Prosternal process slightly incurved
just behind procoxal cavities, then projecting straight towards apex, with clearly emarginated

A New Propsephus from Taiwan

57

Figs. 2-4. Right antenna (2) and aedeagus, dorsal views (3-4) of Propsephus nanshanus sp. nov.,
holotype. Scales: 1 mm (2); 1 mm (3), 0.2 mm (4).

extremity on the underside.

Male genitalia as illustrated in dorsal view (Figs. 3-4 ); median lobe a little longer than
lateral lobes, gradually narrowed towards apex, which is obtusely pointed; each apical portion
of lateral lobe rather triangular, with outer margin clearly and irregularly sinuate and furnished
with many long setae along lateral margin.

Female. Unknown.

Type series. Holotype: d\ Nanshanchi, Nantou Hsien, Taiwan, 12. VIII. 2000, N. Okuda leg.

Etymology. The specific name is derived from the type locality, Nanshanchi.

Notes. This new species is somewhat similar to Propsephus oberthuri (Candeze, 1882)
from Zanzibar, but can be easily distinguished from the later by the following points: 1) body is

58

Hisayuki Arimoto and Sergio Riese

distinctly larger and robuster; 2) dorsal surface is dark brown whereas that of P. oberthuri is
yellowish-brown except for black lateral margins, median longitudinal portion and posterior
angles of pronotum; 3) median longitudinal smooth line on the disk of pronotum is evident
whereas that of P. oberthuri is lacking.

9 to

• Sergio Riese ! £tifz Propsephus JfjCD 1 lit fit - Propsephus

^0, asanas

tl/2 Propsephus 1 ft £rfrft t Propsephus nanshanus L Tffi(Sc L tz .

Propsephus oberthuri Candeze, 1881 <£ %> ,»S, fuKWfSC)

References

Basilewsky, P., 1958. Les Dicrepidiinae du Congo Beige (Coleoptera Elateridae). Entomolgischen
Arbeiten aus Museum G. Frey , 9, Heft 2: 353—477.

Candeze, E., 1859. Monographic des Elaterides 2. Memoires de la Societe royaledes Sciences de Liege,
14: 543 pp., 7 pis.

-, 1882. Elaterides Nouveaux 3. Memoires de la Societe royaledes Sciences de Liege , (2), 9:

ii+117pp.

Hyslop, J. A., 1921. Genotype of the Elaterid beetles of the World. Proceedings of the United states
National Museum , 58: 621-680.

Schenkling, S., 1927. Elateridae H In Junk. W. and S. Schenkling (eds.), Coleopteroum Catalogus , pars
88: 265-636. W. Junk, Berlin.

Schwarz, O., 1902. Neue Elateriden aus dem tropischen Asien, den malaysischen Inseln und den
Inselender Sudsee. Deutsche Entomologische Zeitschift 1902. Heft II: 305-350.

Suzuki, W., 1999. Catalogue of the Family Elateridae (Coleoptera) of Taiwan. Miscellaneous Reports of
the Hiwa Museum for Natural History , (38), 348 pp.

Van Zw aluwenburg , R. H., 1936. The Elaterid beetles of the Philippine Islands. The Philippine Journal
of Science, Manila , 59: 393^132.

(Received February 17, 2005; Accepted March 2, 2005)

Ent. Rev. Japan , 60 (1): 59-62, April 30, 2005

A New Subspecies of Episcaphula matsumurai Chujo
(Coleoptera: Erotylidae) from the Yaeyama Group
in the Southern Ryukyus, Southwestern Japan

Nobuyuki Narukawa
2399 Kida, Suzuka, Mie, 513-0015 Japan
and

Hisashi Ashida

7-4-201, Shimeien, Ibaraki, Osaka, 567-0045 Japan
E-mail: BYD01621 @nifty.ne.jp

Abstract. A new subspecies of Episcaphula matsumurai Chujo, 1941, is described from the
Yaeyama Islands in the southwestern Japan under the name E. matsumurai kaniei Narukawa
et Ashida, ssp. nov.

In 1999, the second author reported the first record of the erotylid genus Episcaphula from
Japan based on a single specimen obtained from the Island of Ishigaki-jima, Okinawa
Prefecture, southwestern Japan, although the real status of the species was not determined
(Ashida, 1999). After that, the first author had an opportunity to examine a series of additional
specimens of that species through the courtesy of Mr. Noboru Kanie. Close examination
revealed that the Japanese population represent a new subspecies of Episcaphula matsumurai
Chujo, 1941, originally described from Taiwan. In this paper, we are going to describe this new
subspecies as a new member of the Japanese erotylid fauna.

Before going further, we wish to express our hearty thanks to Mr. Noboru Kanie (Aichi
Prefecture), Mr. Atsuhiro Nagano (Yamaguchi Prefecture) and Mr. Koji Hosokawa (Aichi
Prefecture) for their kindly offering the valuable materials.

Episcaphula matsumurai kaniei Narukawa et Ashida, subsp. nov.

(Figs. 1-7)

Episcaphula sp.: Ashida, 1999: 11.

Body elongate, oblong-oval, convex on dorsum, about 2.4 times as long as wide, and thin¬
ly clothed with short yellowish brown hairs on the whole surface; general color blackish brown;
elytra decorated with a pair of red anterior bands and of ante-apical spots; anterior band lying
from 2nd interval to shoulder, arcuately emarginate at anterior margin, and undulate at posterior

60

Nobuyuki Narukawa and Hisashi Ashida

margin on 3rd, 5th and 7th striae; the ante-apical spot transversely oval, situated at about apical
fourth of elytron, and occupying from 2nd to 7th striae; scutellum and legs brown.

Head about half as wide as pronotum; punctures sparse, large, and rough; clypeus rather
sharply narrowed towards frontal margin, which is very slightly emarginate at middle; eyes
moderate in size; interocular distance 0.76 times as long as width of head across eyes; maxillary
palpus with terminal segment (Fig. 3) fusiform and about 1.6 times as long as wide; mentum
(Fig. 4) triangular, aciculate apically, with strongly gouged sides in apical two-thirds. Antennae
(Fig. 2) eleven-segmented, with four terminal segments forming a club; 1st segment cylindrical,
wider than long, weakly constricted near apex; 2nd about 1.4 times as long as wide; 3rd about
1.8 times as long as wide, and about 1.3 times as long as 2nd; 4th about 1.6 times as long as
wide; 5th about 1.7 times as long as wide; 6th about 1.4 times as long as wide; 7th about 1.1
times as long as wide; 8th about as long as wide; 9th about 1.2 times as wide as long; 10th about
1.6 times as wide as long; 11th about 1.3 times as wide as long, rounded at apex.

Pronotum about 1.5 times as wide as long, and widest at basal one-third; sides narrowed
towards frontal angles, strongly arcuate at anterior half, and nearly straight at basal half; disk
coarsely and sparsely punctured; anterior margin deeply emarginate and in middle part gently
arched anteriorly; anterior corners roundly projected; posterior comers nearly rectangular.

Elytra elongate, conjointly about 1.5 times as long as wide, widest at basal one-third, and
slightly wider than pronotum; sides weakly arcuate from base to middle, then sharply narrowed
toward apices, with rows eight of seriate punctures, which are relatively finer than those of
pronotum; intervals between rows sparsely and minutely punctured.

Prosternum sparsely punctured; prosternal process (Fig. 5) longer than wide, whose sides
are moderately arcuate at middle and posterior margin is slightly arcuate. Metastemum with
large and rough punctures, and short and sparse hairs.

Median lobe of male genitalia (Figs. 6 and 7) subcylindical; dorsal margin weakly arcuate,
and abruptly declivous in apical ninth, the declivity weakly sinuous; apex pointed in lateral
view, and elongate-ovoid in dorsal view; parameres nearly as long as median lobe; median stmt
long, about 1.2 times as long as median lobe.

Body length: 5.2-5.8 mm; width: 2.1-2.3 mm.

Type series. Holotype: Yonehara, Ishigaki-jima Is., Okinawa Prefecture, 2. V. 2001, N.
Kanie leg. (preserved in the collection of the Osaka Museum of Natural History, Type No.
OMNM-TI-202). Paratypes: 5 exs., same data as for the holotype; 1 ex., same locality as for the
holotype, 9. VI. 1999, N. Kanie leg.; 1?, Mt. Omoto-dake, Ishigaki-jima Is., Okinawa
Prefecture, 3. IV. 1994, A. Nagano leg.; 1 £, same locality, 12. m. 1993, K. Matsumoto leg.;
1 ex., same locality, 29. IV. 2001, K. Hosokawa leg.; 3 exs., Ota, Ishigaki-jima Is., Okinawa
Prefecture, 2. VI. 1988, N. Kanie leg.; 1 ex., same locality, 5. VI. 1988, N. Kanie leg.; 7 exs.,
same locality, 25. VI. 1990, N. Kanie leg.; 1Y, Shiiminato, Iriomote-jima Is., Okinawa
Prefecture, 5~18. IX. 1996, K. Ebi leg. Holotype and four paratypes are preserved in the collec¬
tion of the Osaka Museum of Natural History and other paratypes are in authors' collection.

Distribution. Yaeyama group (Ishigaki-jima Is. and Iriomote-jima Is.) in the southern
Ryukyus, Southwest Japan.

Food-fungus. Unknown.

Etymology. The specific name of the new subspecies is dedicated to Mr. Noboru Kanie.

Notes. The genus Episcaphula Crotch, 1876, is widely distributed in the Southeast Asian
and the Australian Regions and more than 90 species have been described to date (Chujo and

61

A. New

Subspecies of &*«*>*>

matsumurai

Figs

_Dorsal view O*

• • HaR uka*a et ASHiDA, su ^ 5 "° a e d eagas,lateral (6) and

antenna (2), n , .0 m m for t.

dorsal 0) views- Sc

62

Nobuyuki Narukawa and Hisashi Ashida

Chujo, 1988). So far as we are aware, the nominotypical subspecies E. matsumurai matsumurai
is known from only Taiwan and is one of the northernmost species in the genus. Although the
Yaeyama group are closely related to Taiwan in biogeographical viewpoint, this new subspecies
E. matsumurai kaniei from those islands is easily distinguished from the nominotypical sub¬
species by blackish-brown coloration of body and separated ante-apical spots on elytra.

£JH mi • X : + ; n a vJgco

1 frm - +

^ tt + y 3Ay| (ff^) Genus Episcaphula (0 1 ft t L T cf L

A 1999).

tlA Episcaphula matsumurai Chujo, 1941 btltz<D~C,

3 A v Episcaphula matsumurai kaniei Narukawa et Ashida £ L T^p

sb®la. ^ t

(id: 9,

References

Ashida, H., 1999. A new record of erotylid genus Episcaphula from the Island of Ishigakijima.
Nejirebane, Osaka, (83): 11. (In Japanese.)

Chujo, M., 1941. Descriptions of six new erotylid-beetles from Formosa and the Mariana Islands. Mushi ,
Fukuoka, 13: 84-92.

-and Chujo, M. T., 1988. A catalog of the Erotylidae (Insecta, Coleoptera) from the Old World

(excl. the Ethiopian Region). Esakia, Fukuoka, (26): 139-185.

Crotch, G. R., 1876. A revision of the coleopterous family Erotylidae. Cistula Entomology, 1(13):
377-572.

(Received February 24, 2005; Accepted March 4, 2005)

Ent. Rev. Japan , 60 (1): 63-67, April 30, 2005

A New Genus and Species of Coprophilini
(Coleoptera: Staphylinidae: Oxytelinae) from Japan

Yasuhiko Hayashi

Suimeidai 3-1-73, Kawanishi City, Hyogo, 666-0116 Japan

Abstract A new genus and a species of Coprophilini is described from Japan under the name
Coprotrichus kameii.

The members of the tribe Coprophilini are consisting of three genera (Herman, 2001;
Smetana, 2004), which are distributing in Nearctic, Palaearctic, Australian and Neotropical
Regions. Coprophilus Latreille has been known only from Palaearctic Region including
Japan. Recently, I found a peculiar species from Japan, which is well similar in structures of the
gular suture to the genera Homalotrichus Solier from Neotropical and Australian Regions and
Coprostygnus Sharp from New Zealand, but the species is well similar in general appearance to
a Coprophilus species, especially to C. adachii (Y. Watanabe et Y. Shibata) from Japan. After
careful examination, I concluded that the species should be placed in the new genus. In this
paper I am going to describe a new genus for the new species under the name of Coprotrichus
kameii. All type specimens are preserved in the collection of the Osaka Museum of Natural
History, Osaka.

I am very grateful to Dr. Juro Kamei (Akita Pref.) for his kind gift of this interesting material
and to Dr. Katsura Morimoto, the Emeritus Professor of the Kyushu University, Fukuoka, for
his kindness in critically reading the manuscript of this paper.

Coprotrichus gen. nov.

(Figs. 1-4)

Type species. Coprotrichus kameii sp. nov.

Diagnosis. The present new genus is very similar in facies to a Coprophilus- or a
Homalotrichus- species, but it is easily distinguishable from the latter two by absence of distinct
neck. This new genus can be recognized by the combination of the confluent gular suture, the
suborbiculate eyes in lateral view, the single laterosclerite in each side of 3rd to 6th abdominal
segments in appearance in dorsal view, the carinate pro- and mesosternal processes, the contigu¬
ous mesocoxae and 2 rows of spines on dorsum of each tibia.

Description. Body rather small in size, stout, elongate, nearly parallel-sided, flattened
above and well shiny. Elytra and abdomen wider than head and prothorax. Head, pronotum and
elytra with setae scattered sparsely over surface. Abdomen more densely pubescent than fore-

64

Yasuhiko Hayashi

body.

Head (Fig. 2) nearly rounded, uniformly and gently narrowed posteriad, weakly constrict¬
ed at base but without distinct neck, depressed inside supra-antennal ridge and without trans¬
verse groove near posterior margin of eyes on dorsum; epistomal suture absent. Eyes relatively
small, much shorter than postgenae, hemispherically prominent but not extending beyond width
of head and suborbiculate in lateral view. Antennae filiform, moderately long. Labrum short,
strongly transverse, feebly emarginate at apical margin, finely impressed medially, sparsely and
coarsely punctured and fringed with several long and slender setae of various length at apical
margin. Mandibles wide, gently curved ventrad, strongly incurved in apical portion, with a
small tooth near apex and at about the middle of inner margin. Maxillary palpi (Fig. 3) with the
first segment small, nearly as long as wide; second strongly dilated apicad, much longer than the
first; 3rd strongly transverse, much shorter than 2nd, with a few long setae in apical portion; 4th
segment subfusiform, thick, much longer than wide and 3rd, as wide as apex of 3rd in the base.
Labial palpi (Fig. 2) elongate, 1st segment a little wider than long; 2nd much shorter than 1st,
much wider than long; 3rd subconical, much longer than wide and 1st, slightly narrower at the
base than 2nd and subacute at the tip. Ligula wide, weakly emarginate at apical margin.
Paraglossae rather long, extending near to the tip of labial palpi and fringed with sparse short
pubescence in apical portion. Mentum subtrapezoidal, long, nearly twice as wide as long, flat,
coarsely and sparsely punctured, straight at anterior margin, with a long fine seta in antero-later-
al portion. Gular sutures confluent in short length before the middle, divergent anteriad and pos¬
teriad from the confluent portion.

Pronotum obtrapezoidal, narrowed posteriad, weakly sinuate and crenulate at lateral mar¬
gins; disc bearing 3 pairs of depressions, a median ridge and a pair of plaques. Prohypomeron
strongly deflexed and wide. Protergosternal suture present. Procoxal fissure present and open,
and protrochantin well exposed. Postprocoxal lobe (hypomeral projection) present. Prosternal
process carinate in a short distance and extending between coxae.

Scutellum exposed in hind half, tongue-shaped, very shallowly depressed and obsoletely
punctate.

Elytra with epipleural ridge and obscure longitudinal punctate striae. Mesostemal process
elongate, spiniform, short carinate and not strongly extending between mesocoxae, which are
contiguous to each other. Metasternum with a low ridge between mesocoxae. Metasternal
process absent.

Abdomen with 2nd segment not so strongly sclerotized as third, three-fourths as long as
3rd, closely associated with third and apparently not movable on 3rd; 2nd and 3rd sternites
without longitudinal median carina; 3rd to 6th abdominal segment with 2 laterosclerites in each
lateral side but inner laterosclerite not visible in dorsal view; 8th tergite with posterior margin
straight; 8th stemite weakly emarginate at hind margin in male.

Femora with fine pubescence. Tibiae subtrapezoidal in a cross section, bearing two rows
of spines on dorsum and a few longitudinal rows of setae. Mesocoxae not strongly exposed
from mesocoxal cavities and contiguous to each other. Tarsal formula 5-5-5, all segment dis¬
tinct, under side of basal 4 segments with long setae, and 5th segment the longest.

Male genitalia trilobed and symmetrical. Penis subglobular in basal amplitude, strongly
swelling and weakly sclerotized on dorsum, and the apical third abruptly narrowed, parallel¬
sided in the basal half, triangulate in the apical third and acute at the tip. Parameres stout, elon¬
gate and dilated in apical portion.

A New Genus and Species of Coprophilini

65

Figs. 1^4. Coprotrichus kameii sp. nov.-1, Habitus; 2, ventral view of head; 3, right maxilla;

4, ventral view of male genitalia.

Distribution. Japan, Honshu.

Etymology. The generic name is formed by combining parts of the related genera
Coprophilus and Homalotrichus. Both are derived from Greece, and gender of the new name is
masculine.

Discussion. In spite of similar structures of the gular sutures, meso- and metasternal
process of the present new genus it must be placed more closely to the genus Coprophilus
Latreille than to the genus Coprostygnus Sharp. Because Coprostygnus has distinct epistomal
suture, but in the present genus it is absent as in the genera Coprophilus and Homalotrichus.

Coprotrichus kameii sp. nov.

(Figs, l^t)

Body elongate, subparallel-sided, rather flattened above and well shiny; colour reddish
brown to dark reddish brown, elytra, antennae, palpi and legs pale yellow, elytra sometimes
with brownish tinge, and mandibles pitchy. Length: 3.5^4.2 mm.

Head subelliptical, a little longer than wide (in a state well producing in front), gently
roundly narrowed posteriad, gently convex above, vaguely impressed at the top of vertex, rather
deeply depressed inside of antennal ridge, the depression becoming shallower posteriorly, finely
and longitudinally striate in the hind portion, with an obscure and glabrous oblique ridge at the
posterior-most portion; upper surface glabrous in clypeal region, sparsely and coarsely punc¬
tured in frontal region, coarsely and moderately densely punctured in the rest, with striate
microsculpture on postgenae, which is spreaded towards subgenae. Eyes strongly convex, sub-
orbiculate in lateral view and nearly half as long as postgenae (in a state the head well produced

66

Yasuhiko Hayashi

anteriad).

Pronotum a little wider than long (19.5 : 17.0), widest at about anterior fourth, coarsely
crenulate at sides, a little wider than head (19.5 : 17.0); anterior margin weakly bisinuate and
posterior one gently arcuate; from the widest point, sides narrowed posteriad on a straight line;
anterior angles protuberant in front, and posterior ones obtuse; disc gently convex, coarsely and
sparsely punctured, medially with a pair of small depressions just behind the anterior margin, a
pair of shallow longitudinal median depressions in hind two-thirds, and a shallow and large
depression at hind two-thirds of each lateral side which is extending at lateral and basal margin
in the hind half; median ridge between the median depressions glabrous, and a oblique short
glabrous plaque in each lateral side of hind portions of median depressions.

Elytra subtrapezoidal, a little widened posteriad, slightly longer than wide (25.0 : 23.5),
nearly straight at sides except near shoulders and latero-apical angles, which are weakly arcuate,
weakly emarginate at apical margin, with latero-apical angles nearly rounded and inner apical
ones rectangular; surface with 4 obscure longitudinal ridges, which are not reaching base and
apex, and interstices rather densely scattered with small obscure punctures, the punctures some¬
what arranged in a row in lateral 2 interstices.

Abdomen with 3rd to 6th tergites transversely deeply depressed at each base, covered with
distinct fine reticulate microsculpture and sparse and small punctures in a whole, the punctures a
little denser on sternites than on tergites. In female the 8th sternite roundly protuberant behind
and subangulate at the tip.

Legs short and moderately thick; tibiae subtrapezoidal in a cross section, each with 2 rows
of strong spines on dorsal surface, closely setose at under margins and with a longitudinal row
of fine setae on inner and outer surface; each row of spines on protibia consisting of about 7 in
number, those on mesotibiae of about 10, and rather thin and multiple on metatibia.

Penis (Fig. 4) somewhat pear-shaped, abruptly narrowed and strongly curved ventrad in
apical third, which is nearly parallel-sided in the basal half, triangular in distal half and subacute
at the tip; paramere short, curved inward near apex, half length of penis, extending a little
beyond penis and widened inward in distal half.

Type series. Holotype: Mt. Izumigatake, Miyagi Pref., Japan, 9. V. 1965, J. Kamei leg.

Paratypes: 1 <S\ 1 Shizushi, Mizuho-cho, Kyoto Pref., Japan, 13. V. 1982, Y. Hayashi leg.; 1
d\ Ushiroyama, Awakura-mura, Okayama Pref., Japan, 5. V. 1988, A. Watanabe leg.

Etymology. The present species is named in honor of Dr. Juro Kamei, who is an eager
coleopterist in Akita Pref.

# J 1 if JR * fffl. - '>}%

Coprophilini (± 3 b & h /F £ & 1 9 , (i Coprophilus 4r J id 7 X ii *7 i/

Coprophilus adachii (Y. Watanabe et Y. Shibata) T J id 7'^ id 9 vCO/J

— f 7 'J 753^f h HomalotrichusJS,is X ZT~ z. — 'J — y > K1C

Coprostygnusm^X <0 1 #

x. • if ft Coprotrichus kameii ~ ^ 477 # 7 ^ ^ v L np4=r, f£R L tz .

A New Genus and Species of Coprophilini

67

References

Herman, L. H., 1970. Phylogeny and reclassification of the genera of the rove-beetle subfamily
Oxytelinae of the World (Coleoptera, Staphylinidae). Bulletin of the American Museum of Natural
History , 142: 343-454.

-, 2001. The Catalog of Staphylinidae, III, Oxyteline Group. Ibid., 265: 1067-1806.

Smetana, A., 2004: pp. 237-698. In I. Lobl and A. Smetana (ed.): Catalogue of Palaearctic Coleoptera,
Vol. 2. Stenstrup : Apollo Books, 942 pp.

Watanabe, Y. and Y. Shibata, 1961. A revision of the genus Elonium Leach in Japan (Coleoptera,
Staphylinidae). Journal of Agriculture Science, (Tokyo), 7: 43-45.

(Received February 8, 2005; Accepted February 12, 2005)

Ent. Rev. Japan , 60 (1): 68, April 30, 2005

New Records of Staphylinidae from Taiwan, 4

Yasuhiko Hayashi

Suimeidai 3-1-73, Kawanishi City, Hyogo, 666-0116 Japan

1. Philonthus flavipes Kraatz

Specimens examined. 1 ex., Liukuei, 12. VIII. 1972, Y. Maeda leg.; 1 ex., Renting Park, 3. VIII.
1972 (light trap), Y. Maeda leg.; 6 exs.. Is. Lanyu, 24. IV. 1971 (light trap), Y. Hayashi leg.; 1 ex., 2. VI.
1972, Y. Kiyoyama leg.

The species is widely distributed in Southern region of Asia and North Africa.

2. Philonthus fauvelianus Bernhauer

Specimens examined. 1 ex.. Renting Park, 12. VIII. 1969, Y. Maeda leg.; 1 ex., Is. Lanyu, 8. X.
1970, Y. Riyoyama leg.

This species have been known only from Myanmar until now.

3. Philonthus eustilbus Kraatz

Specimens examined. 1 ex., Nanshanchi, 22. IX. 1970, Y. Riyoyama leg. (light trap); 1 ex., ditto, 5.

III. 1970, T. Robayashi leg.; 1 ex., Fungchiifo, 17. VIII. 1969, Y. Maeda leg.; 1 ex., Mt. Yangming, 30.

IV. 1982, T. Ito leg.

This species is widely distributed in Oriental Region. This and P. rutiliventris Sharp, P.
gastralis Sharp (perhaps P. hesperifonnis Cameron, too) are well similar in general appearance
to each other and forming a species group in the genus Philonthus by the structures of the
pronotum and the protarsi. However, they are not true Philonthus-species, because the protarsi
are simply slender, without modified hairs on the undersides as in the genus Gabrius Stephens
or Bisnius Stephens. In consideration of the other characteristics, these species do not belong to
these genera, therefore, I consider they should be transferred to an appropriate genus.

References

Bernhauer, M. and R. Schubert, 1914. Staphylinidae IV. In Junk, W. & S.Schenkling (eds.), Coleo-
pterolum Catalogus, pars 57: 29—408.

Cameron, M., 1932. Coleoptera. Staphylinidae III. In The Fauna of British India including Ceylon and
Burma: xiii+443 pp., 4 pis. Tayler & Francis, London.

Fauvel, A., 1895. Staphylinides nouvaux de Linde et de la Malaisie. Revue d'Entomologie 14: 180-286.
Herman, L. H., 2001. Catalog of Staphylinidae V. Bulletin of the American Museum of Natural History,
265: 2441-3020.

Rraatz, G., 1859. Staphylinen-Fauna von Ostindien, insbesondere der Insel Ceylon. Archiv fur Natur-
geschichte , 25 : 1-196.

(Received February 28, 2005; Accepted March 4, 2005)

Ent. Rev. Japan , 60 (1): 69-74, April 30, 2005

Notes on the Coprophagous Scarab-beetles
(Coleoptera: Scarabaeidae) from Southeast Asia (VI)
— Three New Taxa of the Tribe Coprini from Borneo —

Teruo Ochi

Kohudai 5-21-6, Toyono-cho, Toyono-gun, Osaka, 563-0104 Japan

and

Masahiro Kon

School of Environmental Science, The University of Shiga Prefecture,
Hassaka-cho 2500, Hikone, Shiga, 522-8533 Japan

Abstract Copris ( Copris) poggii sp. nov., C. (C.) gibbulus borneensis subsp. nov. and Micro-
copris fujiokai poringensis subsp. nov. are described from Sabah State, Malaysia.

We had an opportunity to examine Copris specimens collected by T. Kikuta from Kinabalu
National Park, Sabah. As a result, among them, we found a species distinct from any other
species of this genus recorded from Borneo. After a close examination and comparison, we have
concluded that this form is new to science. Thus, we describe a new species of Copris from
Borneo. In addition, we herewith describe two new subspecies of Copris gibbulus Lansberge
and Microcopris fujiokai Ochi et Kon from Borneo, respectively.

Copris {Copris) poggii sp. nov.

(Figs. 1,4-6)

Length: 13.4-19.5 mm; width: 7.0-9.7 mm (n=382).

Body moderate-sized, strongly convex above; dorsal side shining, entirely glabrous; ven¬
tral side also shining; prosternum sparsely clothed with short to long erect yellowish-brown
hairs; mesosternum sparsely clothed with short recumbent yellowish-brown hairs; metasternum
very sparsely clothed with short recumbent yellowish-brown hairs except for glabrous metaster-
nal shield; abdominal sternites clothed with short recumbent yellowish-brown hairs in basal por¬
tion along margin. Color uniformly black, often somewhat reddish; mouth parts, palpi, and
antennae reddish-brown to dark reddish-brown.

Male. Head distinctly transverse; clypeal margin widely emarginated in the middle and
gently rounded on either side, broadly bordered, with a short upturned process at the middle,
which is truncated at apex; apical median portion just below clypeal margin forming a trans¬
verse broad area surrounded by lower and upper (=anterior) margins; the lower margin with

70

Teruo Ochi and Masahiro Kon

three obtuse contiguous teeth in the middle, the median tooth clearly shorter than the outer two;
genae strongly produced laterad, with genal corners right-angled, margin almost straight and
broadly bordered in front, weakly sinuate and finely bordered behind; cephalic horn placed a lit¬
tle behind the middle, short, at most 3 mm or so in length, tapering and pointed distally, clearly
curved backwards; area posterior to the base of cephalic horn almost simple, without a pair of
teeth; vertex transversely and a little shallowly excavated; surface shining, smooth and impunc-
tate in apical half, densely, coarsely and a little shallowly punctate in basal half except for ante¬
rior portion of eye and vertexal excavation almost impunctate.

Pronotum moderately convex, about 1.6-1.7 times as wide as long (n=3), with a distinct
longitudinal impression along midline in basal two-thirds; anterior margin bisinuate, distinctly
bordered, with marginal border widest in the middle and finest laterally; lateral margins gently
rounded in front, slightly sinuous behind, finely bordered; anterior angle rectangular, strongly
produced forwards, with rather sharp corner; basal margin rounded, widely bordered; disc
strongly convex, rather gently declivous in apical third, with the upper edge of the declivity very
obtusely ridged; surface somewhat sparsely covered with coarse and a little shallow punctures,
the punctures becoming very coarse at sides, and also becoming finer on median basal portion
and either side of the declivity.

Elytra strongly convex, about 1.1 times as long as wide (n=3), each with ten striae; 9th and
10th striae confluent in basal third; 1st and 10th, 2nd and 9th, 3rd and 8th, and 4th and 5th
joined at apex respectively; 6th and 7th not distinctly joined at apex; 8th often interrupted
behind the middle; all the striae strongly and rather deeply impressed; strial punctures distinct,
with clearly notching intervals; intervals weakly convex, lightly micro-reticulate though shin¬
ing, sparsely and very finely punctate.

Pygidium weakly convex, shining, densely, coarsely, and a little transversely punctate.
Metasternum with shield almost smooth and impunctate in the middle, sparsely and coarsely
punctate in front and marginal portions. Meso- and metatibiae covered with transverse punc¬
tures, the punctures becoming smaller towards each base though distinct. Protibiae with four
external teeth; the 1st and 2nd teeth contiguous.

Aedeagus rather elongate, about 4.0-4.3 mm (n=3) in total length. Phallobase about
2.1-2.3 mm (n=3) in length, 1.0 mm (n=3) in apical width. Parameres about 1.9-2.0 mm (n=3)
in length; both dorsal lobes forming a short circle, the circle a little opened basad; dorsal mem¬
braneous areas narrow, not developed.

Female. Head with clypeal margin bidentate at the middle, the two teeth small and slightly
reflexed. Pronotum with a slight but distinct depression behind anterior margin.

Type series. Holotype: d\ Headquarter, 1500 m altitude, Mt. Kinabalu, Sabah State,
Malaysia, 3. HI. 1995, T. Kikuta leg. Paratypes: 55 exs.. Poring, Sabah State, Malaysia, 13. IV.
1995, T. Kikuta leg.; 9 exs, ditto, 17. V. 1995; 4 exs., ditto, 700 m alt., 16. V. 1995; 44 exs.,
Headquarter, Kinabalu Park, Sabah State, Malaysia, 3. IV. 1995, T. Kikuta leg.; 1 ex., ditto, 6.

IV. 1995; 6 exs., ditto, 2. V. 1995; 11 exs., ditto, 28. II. 1995: 10 exs., ditto, 3. III. 1995; 51 exs.,
ditto, 12. X. 1997; 21 exs., Tahubang, Sabah State, 19. IV. 1995, T. Kikuta leg.; 32 exs., ditto,
20. IV. 1995; 10 exs., ditto, 24. n. 1995; 2 exs., ditto, 24.1. 1995; 115 exs., Sayap, nr. Kinabalu,
Sabah State, Malaysia, 25. III. 1995, T. Kikuta leg.; 10 exs., ditto, 12. V. 1995; 2 exs., ditto, 12.

V. 1995.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Three New Taxa of Coprini from Borneo

71

Figs. 1-3. 1, Copris ( Copris) poggii sp. nov., male, habitus, dorsal view; 2, C. (C.) gibbulus
borneensis subsp. nov., male, habitus, dorsal view; 3, Microcopris fujiokai poringensis subsp.
nov., male, habitus, dorsal view.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. This species is named in honor of Dr. Roberto Poggi, Museo Civico di Storia
Naturale di Genova (Giacomo Doria), who has been giving the first author invaluable help for
his researches.

Notes. The present new species is closely related to Copris ( Copris) agnus Sharp, from
Malay Peninsula and Borneo, but can be distinguished from the latter by the following charac¬
teristics: 1) pronotum declivous in front, with a distinct depression at the middle of the declivity
in both sexes, whereas in C. agnus , it is simple without a distinct depression in front; 2) meso-
and metatibiae with each ventral side clearly punctate near basal part, whereas in C. agnus , they
are almost impunctate near basal part; 3) in the male, head with clypeal margin bearing a short
upturned process instead of being a pair of strongly reflexed teeth at the middle of clypeal mar¬
gin; 4) in the male, head with cephalic horn short but clearly curved backwards, whereas in C.
agnus , it is very short, conical, and vertical; 5) in the male genitalia, the parameres with both
dorsal lobes forming a short circle, and dorsal membraneous areas are very short and narrow,
whereas in C. agnus , the parameres with the dorsal lobes form an elongate circle and the dorsal
membraneous areas are long and wide (cf. Fig. 7).

Copris ( Copris) gibbulus borneensis subsp. nov.

(Fig. 2)

The present new subspecies differs from the nominotypical one from Borneo as follows:
1) body larger (11.0-14.0 mm), whereas in the nominotypical subspecies, the body length is a
little smaller (9.0-12.0 mm); 2) head with clypeus distinctly punctate on the posterior portion,

72

Teruo Ochi and Masahiro Kon

Figs. 4-7. Copris ( Copris ) spp.-4, C. (C.) poggii sp. nov., male, right protibia; 5, aedeagus,

lateral view; 6, aedeagus, dorsal view.-7, C. (C.) agnus Sharp, parameres, dorsal

view.

whereas in the nominotypical subspecies, it is almost impunctate or provided with ill-defined
small punctures; 3) pronotum with anterior angle less produced forwards, with corner rounded
in small male and female, whereas in the nominotypical subspecies, the anterior angle of prono¬
tum is more strongly produced forwards with the corner distinctly sharp in small male and
female; 4) elytra with intervals distinctly convex and very finely punctate to almost impunctate
instead of being weakly convex and finely punctate; 5) elytra with striae clearly wider.

Length: 11.0-14.4 mm; width: 5.1-7.2 mm (n=6).

Type series. Holotype. <?, Keningau, Sabah State, Malaysia, V. 1997. Paratypes: 2-?--?-,
the same data as the holotype; 1 ?, Crocker Range, Sabah State, Malaysia, 12. VI. 1994; <?,

Mt. Bawang, Kalimantan, Indonesia, VII. 1991.

Type depository. The holotype is at present preserved in the collection of the National
Science Museum (Natural History), Tokyo.

Distribution. Sabah State, Malaysia; West Kalimantan, Indonesia (Northern and Western

Three New Taxa of Coprini from Borneo

73

Borneo).

Etymology. The subspecies is named after Borneo.

Microcopris fujiokai poringensis subsp. nov.

(Fig. 3)

The present new subspecies differs from the nominotypical one from Kalimantan,
Indonesia, West Borneo as follows: 1) body size clearly larger (9.5-11.2 mm), whereas in the
nominotypical subspecies, the body is relatively smaller (7.7-9.9 mm); 2) elytra with interval
mostly opaque, rarely a little lustrous, and distinctly micro-granulose to micro-reticulate instead
of being strongly shining; 2) body more distinctly suffused with purplish luster, especially on
elytra, whereas in the nominotypical subspecies, the body is usually suffused with slight pur¬
plish luster on head and pronotum, without metallic luster on elytra; 3) head with median tuber¬
cle on frons stronger; 4) head and pronotum less coarsely and more sparsely punctate.

Length: 9.5-11.2 mm; width: 5.0-5.9 mm (n=14).

Type series. Holotype: <?, Poring, 1200 m alt., Sabah State, Malaysia, 13. IV. 1995, T.
Kikuta leg. Paratypes: 8 exs., the same data as the holotype; 9 exs., 16.1. 1997, the same locali¬
ty as the holotype.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. The subspecies is named after a place name, Poring, Sabah State.

Acknowledgments

We wish to express our cordial thanks to Mr. T. Kikuta for giving us the opportunities of
examining invaluable specimens. This study was supported in part by a Grant-in-Aid from the
Japan Society for the Promotion of Science (No. 14405013).

51 ft)

VM * & Hit : A (iM$K) — 401/^ tMVd 3 ir u if

3 Sr® — . - 401/4-4-ji 3 LT, Vd 3^3 #'4-SO 1 Sr®

Copris ( Copris ) poggii sp. nov. t , |W|fl|<7)fji C. (C.) gibbulus Lansberge CO 1 SfjEfi C. (C.)
gibbulus borneensis subsp. nov. jo <£ t ?d u jr a |Ofl Microcopris fujiokai Ochi et
Kon (D 1 firStfi M. fujiokai poringensis subsp. nov. i 401/ 4-jtSM* L tz.

References

Balthasar, V., 1963. Monographic der Scarabaeidae und Aphodiidae der palaearktischen und orientalis-
che Region, 1: 1-391. Prag.

Boucomont, A., 1914. Les Coprophages de l’Archipel Malais. Annales de la Societe. Entomologique de
France , 83: 238-350.

74

Teruo Ochi and Masahiro Kon

Lansberge, J. W. v., 1886. Les Coprides de la Malaisie. Tijdschrift voor Entomologie, 29: 1-25.

Ochi, T. and M. Kon, 1994. Dung beetles (Coleoptera, Scarabaeoidea) collected from Sabah, Borneo (1).
Elytra, Tokyo, 22: 281-298.

-and-, 1996. Studies on the coprophagous scarab beetles from East Asia. IV (Coleoptera,

Scarabaeidae). Giornale italiano di Entomologia , 8: 17-28.

Sharp, D., 1875. IV. Description of some new genera and species of Scarabaeidae from tropical Asia and
Malaysia, Part I. Coleopterologische Hejfte , 13: 33-54.

(Received March 18, 2005; Accepted April 11, 2005)

Em. Rev. Japan , 60 (1): 75-82, April 30, 2005

Notes on the Coprophagous Scarab-beetles
(Coleoptera: Scarabaeidae) from Southeast Asia (VII)

— Three New Species of Onthophagus ( Phanaeomorphus ) from Borneo —

Teruo Ochi

Kohudai 5-21-6, Toyono-cho, Toyono-gun, Osaka, 563-0104 Japan

and

Masahiro Kon

School of Environmental Science, The University of Shiga Prefecture,
Hassaka-cho 2500, Hikone, Shiga, 522-8533 Japan

Abstract Three new species of the genus Onthophagus (Phanaeomorphus ) are described
from Borneo under the names of O. ( P .) quasijohkii sp. nov., O. ( P.) maryatiae sp. nov. and
O. ( P .) cjuasitagal sp. nov.

In 1935, Balthasar erected the subgenus Phanaeomorphus in the genus Onthophagus
Latreille based on a Chinese species, Onthophagus ( Phanaeomorphus ) sycophanta Fair-
maire. As far as we are aware, ten or so species are known from Asia. From Borneo, we record¬
ed two species belonging to this subgenus, O. ( P .) bangueyensis Boucomont and O. (P.) johkii
Ochi et Kon in 1994.

When we examined Onthophagus specimens collected by T. Kikuta from Mt. Kinabalu, we
found three undescribed species of the subgenus Phanaeomorphus among them. Thus, we
describe three new species of Phanaeomorphus from Borneo.

Onthophagus ( Phanaeomorphus ) quasijohkii sp. nov.

(Figs. 1,4-6)

Length: 6.6-8.8 mm; width: 3.5-4.2 mm (n=388).

Body small-sized, rather elongate-oval, strongly convex; dorsal side mat, a little densely
clothed with short suberect yellowish-white hairs, except for glabrous head. Color blackish-
brown to grayish-black, with a weak greenish tinge; mouth organs, palpi, antennal foot-stalks,
and legs somewhat reddish; club segments of antenna dark yellowish-brown.

Male. Head almost simple, polygonal in outline; clypeus strongly produced forwards as a
reflexed rounded subtriangular lobe at the middle, the lobe about 0.5 mm in length in large
males; in small males, the lobe reduced to a blunt small tooth; clypeo-frontal suture completely
effaced, clypeo-genal suture fine and not carinate though barely perceptible; genae produced lat-

76

Teruo Ochi and Masahiro Kon

erad, with genal corner obtusely angulate at the middle; vertex slightly raised at the middle in
the posterior most part; surface micro-granulose except for a little shining clypeus, and closely
covered with coarse punctures, the punctures uneven and weakly wrinkled at clypeus, and
changing into more crowded annular ones towards vertex.

Pronotum strongly convex, about 1.30-1.37 times as wide as long (n=5), with an obsolete
longitudinal impression along midline in basal half; anterior margin weakly bisinuate, distinctly
bordered; lateral margins gently rounded in front, very weakly sinuate behind, finely bordered;
anterior angles strongly produced forwards, with apices slightly expanded outwards; posterior
angles obtuse; basal margin gently rounded, or very obtusely angulate at the middle, finely bor¬
dered; disc declivous towards both anterior angles in front, leaving the posterior part triangular¬
ly elevated; upper edge of the declivity sharply carinate on both sides, the carina becoming
gradually obsolete towards the median angle, which is very blunt; in small males, the triangular
part of disc almost reduced; surface distinctly micro-granulose, and densely covered with rather
coarse ocellate shallow punctures, the punctures becoming denser and larger towards sides and
base.

Elytra about 1.13-1.21 times as wide as long (n=3), with eight striae including one along
epipleural margin; each stria rather widely, a little shallowly, and strongly impressed, with fine
ridge on both sides throughout, also becoming fairly deep at the 5th to 8th striae; 7th not dis¬
tinctly curved; strial punctures obviously transverse, with clearly notching intervals; each punc¬
ture separated into two round bottoms; intervals almost flat, strongly micro-granulose, and
somewhat sparsely covered with small granules or asperate-punctures.

Pygidium well convex, carinate at base, weakly micro-granulose, densely covered with
coarse ocellate round punctures. Meso- and metatibiae a little densely covered with transverse
coarse punctures throughout on each ventral side. Protibiae rather elongate, with four external
teeth; 1st and 2nd teeth contiguous, 3rd a little separated from the 2nd; terminal spur finger-like,
slightly decurved, pointed apically.

Aedeagus clearly larger than those in the related species. Phallobase about 1.5-1.6 mm in
length (n=3), about 0.7 mm in apical width (n=3). Parameres about 0.7-0.8 mm in length, with
apices a little expanded and toothed outwards in dorsal view.

Female. Head widely emarginated in the middle, with the median portion strongly pro¬
duced as a reflexed short process, the process forked into two sharp apices distally; surface more
densely punctate than in male. Pronotum also declivous towards both anterior angles, though the
upper edge of the declivity more obtuse and not sharply carinate on both sides; surface more
densely and coarsely punctate than in male.

Type series. Holotype: c?, Headquarter, Kinabalu Park, Sabah State, Malaysia, 12. II.
1995, T. Kikuta leg. Allotype: the same data as for the holotype. Paratypes: 15 exs., the

sama data as for the holotype; 10 exs., ditto, X. 1994, T. Kikuta leg.; 27 exs., ditto, 2. II. 1995;
10 exs., ditto, III. 1995. T. Kikuta leg.; 18 exs., ditto, 3. IV. 1995; 4 exs., ditto, 28. II. 1995; 11
exs., ditto, 3. III. 1995; 30 exs., ditto, 4. III. 1995; 4 exs., ditto, 1. V. 1995; 1 ex., Liwagu, Sabah
State, Malaysia, X. 1994, T. Kikuta leg.; 1 ex., ditto, VI. 1995, T. Kikuta leg.; 2 exs., ditto, 4.
VI. 1995; 1 ex., ditto, 6. IV. 1995; 1 ex., 9. IV. 1995; 78 exs., Tahubang, Sabah State, Malaysia,
24. I. 1995, T. Kikuta leg.; 37 exs., ditto, 20. IV. 1995; 2 exs., ditto, 19. IV. 1995; 18 exs.,
ditto, 19. IV. 1995; 93 exs., Sayap, nr. Kinabalu, Sabah State, Malaysia, 12. V. 1995, T. Kikuta
leg.; 8 exs., ditto, 25. III. 1995; 7 exs., ditto, 8. XI. 1995; 6 exs., ditto, 11. V. 1995; 1 ex.,
Poring, Sabah State, Malaysia, 13. IV. 1995, T. Kikuta leg.; 1 ex., ditto, 17. V. 1995.

Three New Species of Onthophagus from Borneo

77

Figs. 1-3. Onthophagus (Phanaeomorphus) spp., male, habitus, dorsal views.-1, O. ( P .) quasijohkii sp.

nov.; 2, O. ( P .) maryatiae sp. nov.; 3, O. (P.) quasitagal sp. nov.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. The specific name means that the present new species is similar to Ontho¬
phagus ( Phanaeomorphus) johkii Ocht et Kon.

Notes. The present new species is closely related to Onthophagus johkii Ochi et Kon from
Borneo, but can be distinguished from the latter by the following characteristics: 1) punctures
on basal portion of pronotum coarser and clearly shallower, whereas in O. johkii , they are fairly
coarse and rather deep: 2) meso- and metatibiae with each ventral side densely covered with
transverses coarse punctures throughout, whereas in O. johkii, it is sparsely punctate on basal
half, not dense throughout; 3) elytra with stria rather wide instead of being clearly wide; 4) in
the male, frons entirely flat, whereas in O. johkii , it is obtusely and transversely raised; 5) in the
female, head with the median process forked into two sharp apices distally; 6) male genitalia
larger and different in shape.

Onthophagus (Phanaeomorphus) maryatiae sp. nov.

(Figs. 2, 7)

Length: 7.7-9.0 mm; width: 4.1-5.0 mm (n=4).

Body a little larger in size, oval, strongly convex; dorsal side shining, almost glabrous;
ventral side also shining, partly clothed with reddish-brown hairs. Color uniformly black to red¬
dish-brown; mouth organs, palpi, legs more or less reddish; antennae reddish-brown with club
segments dark yellowish-brown.

Male. Head subpentagonal; clypeus strongly and sub-triangularly produced forwards, with

78

Teruo Ochi and Masahiro Kon

apex upturned as a small and obtusely bidentate lobe at the middle; genae strongly protrudent
laterad, with genal corner rounded; clypeal suture clearly and straightly carinate at frontal sec¬
tion, not carinate at genal sections; vertex with a short transverse carina at the middle in the pos-
teriormost part; surface shining in front, weakly micro-granulose behind, densely, coarsely, and
a little vaguely punctate except for clypeus where the punctures are becoming finer, sparser, and
transversely wrinkled.

Pronotum strongly convex, about 1.34-1.39 times as wide as long (n=3), with an obsolete
longitudinal impression along midline in basal third; anterior margin bisinuate, rather thickly
bordered; lateral margins strongly rounded at the middle, sinuate behind, thinly bordered; ante¬
rior angles strongly projected anteriad, subrectangular, with corner rounded and a little expand¬
ed outward; posterior angles obtuse; basal margin obtusely angulate in the middle, finely bor¬
dered; disc declivous towards both anterior angles in front, leaving the posterior part widely and
triangularly elevated; the upper edge of the declivity briefly carinate on both sides, the carina
gradually becoming obsolete towards the obtuse median angle; surface distinctly shining on tri¬
angular posterior portion, weakly micro-granulose at anterior declivities, and regularly with
sparse fine punctures intermixed with rather sparse, coarse, and annular ones, and both punc¬
tures becoming denser and larger at the declivities.

Elytra about 1.28-1.39 times as wide as long (n=3), strongly convex, with eight striae
including one along epipleural margin; each stria rather finely, shallowly impressed, with fine
ridge on both sides throughout; strial punctures obviously transverse, with slightly notching
intervals; each puncture separated into two bottoms; 7th stria almost parallel to 6th; intervals
weakly convex, slightly micro-granulose though shining, rather sparsely covered with ill-
defined small punctures.

Pygidium slightly convex near apex, carinate at base, very uneven, weakly micro-granu¬
lose, sparsely covered with a little transverse ocellate punctures. Meso- and metatibiae shining,
smooth, very sparsely and finely punctate on each ventral side, with a few coarse and scattered
punctures. Protibiae rather elongate, with four external sharp teeth; terminal spur ordinarily
sharp, well decurved.

Aedeagus rather small. Phallobase about 1.2 mm in length (n=l), about 0.6 mm in apical
width (n=l). Parameres about 0.7 mm (n=l), each with ventral tooth near apex from lateral
view; from dorsal view, each apical portion a little expanded outwards.

Female . Head with clypeus less produced forwards and less reflexed; front-clypeal suture
gently and evenly curved; clypeus more weakly rugose. Pronotum with triangular part of disc a
little obtusely ridged in front. Elytra less lustrous. Protibiae with four external teeth stronger.

Type series. Holotype: <?, Headquarter (Liwagu), 1450 m, Kinabalu Park, Sabah State,
Malaysia, X. 1994, T. Kikuta leg. Paratypes: 1 <?, the same data as the holotype; 1 c?, ditto, 2.
III. 1995, T. Kikuta leg.; 1 £, ditto, 29. X. 1998.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Etymology. This species is named in honor of Prof. Maryati Mohamed, University Malay¬
sia Sabah, who has been giving us invaluable help for our researches in Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Notes. The present new species is closely related to Onthophagus (Phanaeomorphus )
tagal Boucomont from the Philippines, but can be distinguished from the latter by the follow¬
ing characteristics: 1) body distinctly larger; 2) pronotum with lateral margins not strongly

Three New Species of Onthophagus from Borneo

Figs. 4-9. Onthophagus (Phanaeomorphus) spp.-4, O. (P.) quasijohkii sp. nov., aedeagus. dorsal

and lateral views; 5, male, head and anterior part of pronotum, dorsal view; 6, female, head, dor¬
sal view.-7, O. ( P .) maryatiae sp. nov., aedeagus, dorsal and lateral views.-8, O. ( P .)

quasitagal sp. nov., aedeagus, dorsal and lateral views.-9, O. ( P .) tagal Boucomont, aedea¬

gus, lateral view.

80

Teruo Ochi and Masahiro Kon

rounded in the middle, whereas in O. tagal, it is strongly rounded in the middle; 3) in the male,
head with vertex slightly raised at the middle, whereas in O. tcigal, it is strongly raised at the
middle; 4) in the male, aedeagus clearly different in shape (cf. Fig. 9).

Onthophagus ( Phanaeomorphus ) quasitagal sp. nov.

(Figs. 3, 8)

Length: 5.4-7.9 mm; width: 3.1^4.3 mm (n=364).

Body moderate-sized, oval, strongly convex above; dorsal side fairly shining, entirely
glabrous; ventral side also shining, partly clothed with reddish-brown hairs. Color uniformly
black to reddish-brown, frequently with a slight purplish tinge on head; mouth organs, palpi,
legs more or less reddish; antennae reddish-brown with club segments dark yellowish-brown.

Male. Head sub-pentagonal; clypeus strongly and subtriangularly produced anteriad, with
apex reflexed as a small lobe at the middle, the lobe almost truncated distally in large males; in
small males, the lobe reduced to short bidentate teeth; genae strongly protrudent laterad, with
genal corner obtusely angulate; clypeal suture with frontal section completely effaced, genal
section not carinate; vertex with a short transverse carina at the middle in the posteriormost part,
the carina not strongly raised; surface shining, fairly densely and strongly punctate, the punc¬
tures becoming more and transversely wrinkled.

Pronotum strongly convex, about 1.40-1.50 times as wide as long (n=5); median longitu¬
dinal impression not distinct; anterior margin bisinuate, rather thickly bordered; lateral margins
strongly rounded at the middle, almost straight in front, sinuate behind, and thinly bordered;
anterior angles strongly produced forwards, sub-rectangular, with corner rounded and a little
expanded outwards; posterior angles obtuse; basal margin obtusely angulate in the middle, thin¬
ly bordered; disc declivous towards both anterior angles in front, leaving the posterior part more
widely and triangularly elevated than in the proceeding species, though the upper edge of the
declivities more obtusely carinate; median angle of the triangular disc very obtuse; in smaller
males, the triangular disc becoming almost simple; surface very shining on triangular posterior
portion, weakly micro-granulose at anterior declivities, rather sparsely covered with shallow
annular punctures, the interspaces between punctures bearing fine and sparse punctures, both
punctures becoming denser and stronger towards sides.

Elytra strongly convex, about 1.21-1.36 times as wide as long (n=5); disc with eight striae
including one along epipleural margin; each stria rather shallowly impressed with fine ridge on
both sides throughout; strial punctures a little transverse, with slightly notching intervals; each
puncture separated into tow round buttoms; 7th stria almost parallel to the 6th; intervals almost
flat, shining, rather sparsely covered with small punctures.

Pygidium gently convex near apex, carinate at base, a little uneven, shining, moderately
densely covered with transverse ocellate punctures. Meso- and metatibiae with each ventral side
shining smooth, sparsely and finely punctate. Protibiae elongate, with four external sharp teeth;
terminal spur ordinarily sharp, well decurved.

Aedeagus rather slender. Phallobase about 1.1-1.5 mm in length (n=3), about 0.5-0.7 mm
in apical width (n=3). Parameres about 0.7-0.8 mm (n=l), each with ventral small sharp tooth
near apex from lateral view; from dorsal view, each apex distinctly produced outwards.

Female. Head with clypeal margin rather broadly truncated or slightly emarginated at the

Three New Species of Onthophagus from Borneo

81

middle. Pronotum very weakly declivous towards both anterior angles, the triangular disc barely
perceptible. Protibiae with four external teeth stronger; terminal spur more sharply pointed.

Type series. Holotype: c?\ Sayap, nr. Kinabalu, Sabah State, Malaysia, 8. XI. 1994, T.
Kikuta leg. Paratypes: 2 exs., Poring, Sabah State, Malaysia, 13. IV. 1995, T. Kikuta leg.; 52
exs., Headquarter, Kinabalu Park, Sabah State, Malaysia, X. 1994, T. Kikuta leg.; 32 exs.,
ditto, 2. II. 1995; 19 exs., ditto, 12. II. 1995; 1 ex., ditto, III. 1995, T. Kikuta leg.; 27 exs., ditto,
3. IV. 1995; 4 exs., ditto, 2. V. 1995; 11 exs., ditto, 2. III. 1995; 7 exs., ditto, 4. III. 1995; 92
exs., Liwagu, Sabah State, Malaysia, X. 1994, T. Kikuta leg.; 1 ex., ditto, VI. 1995; 2 exs.,
ditto, 4. VI. 1995; 4 exs., Tahubang, Sabah State, Malaysia, 20. IV. 1995, T. Kikuta leg.; 4
exs., ditto, 19. IV. 1995; 10 exs., ditto, 24. I. 1995; 17 exs., Sayap, nr. Kinabalu, Sabah State,
Malaysia, 12. V. 1995, T. Kikuta leg.; 7 exs., ditto, 25. III. 1995; 8 exs., ditto, 7. XI. 1995; 29
exs., ditto, 8. XI. 1995; 3 exs., ditto, 11. V. 1995.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. The specific name means that the present new species is similar to Ontho¬
phagus (Phanaeomorphus) tagal Boucomont.

Notes. The present new species is closely related to Onthophagus ( Phanaeomorphus)
tagal Boucomont from the Philippines, but can be distinguished from the latter by the follow¬
ing characteristics: 1) body distinctly larger; 2) head without a transverse carina on frons in both
sexes, whereas in O. tagal , head with a distinct transverse carina on frons in the small male and
female; 3) head with vertex transversely and rather weakly raised at the middle, whereas in O.
tagal it is strongly raised at the middle; 4) in the male, aedeagus clearly different in shape (cf.
Fig. 9).

Acknowledgments

We wish to express our cordial thanks to Mr. T. Kikuta for giving us the opportunities of
examining invaluable specimens. This study was supported in part by a Grant-in-Aid from the
Japan Society for the Promotion of Science (No. 14405013).

5 ? ft)

®03Sffl—. - It, 1ES,

Phanaeomorphus^MD 3 frS, Onthophagus ( Phanaeomorphus) quasijohkii sp. nov., O. (P.)
maryatiae sp. nov., O. (P.) quasitagal sp. nov. ffi® L tz .

References

Balthasar, V., 1935. Onthophagus -Arten Chinas, Japans, und der angrenzenden Lander. Folia
Zoologica et Hydrobiologica, 8: 303-353.

-, 1963. Monographic der Scarabaeidae und Aphodiidae der palaearktischen und orientalische

Region, 2: 1-628. Prag.

82

Teruo Ochi and Masahiro Kon

Boucomont, A., 1914. Les Coprophages de l'Archipel Malais. Annales de la Societe Entomologique de
France , 83: 238-350.

Boucomont, A., 1914. Onthophagus asiatiques nouveaux ou peu connus. Annali Museo Civico di
Genova , 46: 210-243.

-, 1924. Les Onthophagus (Coleoptera, Scarabaeidae) des lies Philippines. Philippine Journal of

Science , 24: 669-681.

Ochi, T. and M. Kon, 1994. Dung beetles (Coleoptera, Scarabaeoidea) collected from Sabah, Borneo (1).
Elytra, Tokyo , 22: 281-298.

(Received March 18. 2005; Accepted April 11, 2005)

Ent. Rev. Japan, 60 (1): 83-98, April 30, 2005

Notes on the Coprophagous Scarab-beetles
(Coleoptera: Scarabaeidae) from Southeast Asia (VIII)
— Six New Species of Onthophagus ( Parascatonomus ) and
a New Subspecies of O. (P.) katoi from Borneo —

Teruo Ochi

Kohudai 5-21-6, Toyono-cho, Toyono-gun, Osaka, 563-0104 Japan

and

Masahiro Kon

School of Environmental Science, The University of Shiga Prefecture,
Hassaka-cho 2500, Hikone, Shiga, 522-8533 Japan

Abstract Six new species of Onthophagus (Parascatonomus ) are described from Borneo
under the names of O. (P.) riekoae sp. nov., O. (P.) liewi sp. nov., O. ( P .) kikutai sp. nov., O.
(P.) anitidus sp. nov., O. (P.) sayapensis sp. nov., and O. (P.) gunsalami sp. nov. In addition,
a new subspecies of O. ( P .) katoi is also described from Borneo under the name of O. (P.)
katoi poringensis subsp. nov.

Paulian (1932) originally described Parascatonomus as a genus in the tribe Onthophagini
Afterward, and he (1945) placed it as a subgenus of the genus Onthophagus Latreille. Later,
Ochi and Araya (1992) revised the definition of this subgenus. According to their definition,
Kon et al. (2000) recorded 12 species of Parascatonomus from Borneo.

When we examined Bornean specimens of Parascatonomus in our hands, we found six
undescribed species among them. In addition, we found that the Bornean specimens that had
previously been identified with O. (P.) pilularius Lansberge were distinct from the topotype of
O. (P.) pilularius from Java. After a close examination, this Bornean form was identified with
O. (P.) katoi Ochi et Araya, although it is slightly different from the holotype of O. (P.) katoi
from the Philippines. Thus, we describe six new species of Parascatonomus and a new sub¬
species of O. (P.) katoi from Borneo.

84

Teruo Ochi and Masahiro Kon

Onthophagus ( Parascatonomus ) riekoae sp. nov.

(Fig. 1)

Length: 13.2 mm; width: 6.3 mm (n=l).

Female. Body large-sized, rather depressed dorsally, and shallowly constricted between
pronotum and elytra; dorsal surface shining, sparsely clothed with short yellowish hairs; ventral
surface also shining, clothed with similar hairs as those on dorsum. Color almost black, partly
with slight purplish luster, especially on pygidium and abdominal sternites.

Head subpentagonal; clypeus a little produced forwards, with anterior margin parabolic in
outline, slightly reflexed at the middle; clypeo-frontal suture completely effaced though obtuse¬
ly raised at the middle; genae well produced laterad, with genal angle a little wider than a right
angle; vertex with a well curved transverse obtuse carina whose both sides are strongly raised,
and with a round shallow concavity in front of the carina at the middle; surface transversely
rugose in apical half, the rugosities changed into round granules in basal half. Antennae short
and compact; scape fairly short, invisible from dorsal aspect; the 1st segment large, a little
longer than O. discedens, enclosing basal portion of the 2nd; 4th to 6th closely joined, not
strongly broadened distally as in O. discedens', antennal club small.

Pronotum somewhat convex though a little depressed dorsally, about 1.4 times as wide as
long (n=l), with a slight impression in basal two-thirds along midline; anterior margin emar-
ginated, finely bordered; lateral margins gently and evenly rounded in front, not distinctly sinu¬
ate behind, finely bordered throughout; anterior angle rounded though well protrudent forwards;
posterior angle rounded; base very obtusely angulate and slightly raised at the middle, not dis¬
tinctly bordered; disc declivous in anterior third, with the declivity weakly and obtusely raised
along midline and a little depressed laterally; surface shining, evenly and rather sparsely cov¬
ered with shallow setiferous punctures, the punctures small at the middle, and changing into a
round granules anteriorly and towards sides.

Elytra rather elongate and a little depressed dorsally, about 1.2 times as wide as long
(n=l), with eight striae including one along epipleural margin; each stria distinctly impressed
though a little shallow, clearly wider than in O. discedens ; 7th stria clearly curved near base;
intervals flat, shining, evenly and somewhat sparsely covered with small setiferous punctures.

Pygidium carinate at base, densely and strongly punctate. Prosternum with anterior angle
broadly and not so deeply excavated on the ventral side. Metastemum with antero-median por¬
tion very slightly raised and briefly keeled along midline a little behind the elevation. Protibiae
clearly broad and short, with four lateral blunt teeth; 4th tooth very obtuse. Meso- and metatarsi
with 2nd to 5th segments clearly short.

Male. Unknown.

Type specimen. Holotype: Mt. Bawang, W. Kalimantan, VIII. 1993, N. Nishikawa leg.

Type depository. The holotype is preserved in the Osaka Museum of Natural History.

Distribution. West Kalimantan, Indonesia (Southwestern Borneo).

Etymology. This species is named in honor of Professor Rieko Muramoto.

Notes. The present new species is somewhat related to Onthophagus discedens Sharp
from the Sunda Islands, but can be distinguished from the latter by the following characters; 1)
head with anterior margin simple; 2) head with vertex strongly carinate instead of being simple;
3) pronotum with a declivity in front, whereas in O. discedens , it is entirely simple; 4) meso-
and metatarsi clearly shorter.

Six New Species and a New Subspecies of Onthophagus from Borneo

85

Figs. 1-7. Onthophagus (Parascatonomus) spp., habitus, dorsal views.-1, O. ( P .) riekoae sp. nov., female;

2, O. ( P .) liewi sp. nov., male; 3, O. ( P .) kikutai sp. nov., male; 4, O. ( P .) anitidus sp. nov., male; 5, O. (P.)
gunsalami sp. nov., male; 6, O. (P.) sayapensis sp. nov., male; 7, O. (P.) katoi poringensis subsp. nov.,
male.

86

Teruo Ochi and Masahiro Kon

Figs. 8-9. Onthophcigus ( Parascatonomus) spp., male clypeal margin, dorsal views.-8, O. (P.)

liewi sp. nov.; 9, O. (P.) sarawacus Harold.

Onthophagus ( Parascatonomus ) liewi sp. nov.

(Figs. 2, 8, 10-11)

Length: 10.4-14.4 mm; width: 6.2-8.4 mm (n=248).

Body large, oblong-oval, strongly convex; dorsal side mat, almost glabrous, with prono-
tum often bearing several semirecumbent long hairs on each side near base; ventral side slightly
shining, partly and densely clothed with reddish hairs. Color black; mouth organs, palpi and legs
more or less reddish; antennae reddish-brown with club segments yellowish-brown on apical
half.

Male. Head about 1.23 to 1.29 times as wide as long (n=10); clypeus strongly produced
forwards as a reflexed, slightly prolonged and rounded tooth in the middle, with basal sides of
the tooth shallowly but clearly incised; clypeal margin except for the median tooth reflexed and
broadly bordered; clypeo-frontal suture barely perceptible or almost effaced; genae well pro¬
duced laterad, obtusely angulate at the middle; vertex with a transverse obtuse carina a little
before posterior margin, which is only slightly postcurved, clearly raised; surface densely and
transversely rugose and strongly punctate, the rugosities partly changing into granules. Anten¬
nae short and compact; scape short, invisible from dorsal aspect; club segments successively
diminishing the size in breadth and length.

Pronotum strongly convex, about 1.40 to 1.50 times as wide as long (n=10), with an obso¬
lete longitudinal impression along midline; anterior margin bisinuate for receiving the posterior

Six New Species and a New Subspecies of Onthophagus from Borneo

87

Figs. 10-12. Onthophagus (Parascatonomus ) spp.-10, O. ( P .) liewi sp. nov., male, head and

pronotum, dorsal view; 11, aedeagus, dorsal and lateral views.- 12, O. ( P .) sarawacus

Harold, aedeagus, dorsal and lateral views.

portion of head, finely bordered; lateral margins evenly and rather strongly rounded in front,
sinuate behind, with distinct marginal line; basal margin slightly produced posteriad and angu-
late at the middle, with marginal line bordered at the middle and unbordered laterally; anterior
angles bluntly subangulate; posterior angles obtuse; disc somewhat gently declivous in anterior
third, with the declivity slightly depressed in major males; whereas in minor males, the declivity
becoming smaller and indistinct; surface strongly micro-granulose, sparsely and finely punctate
on disc, the punctures gradually changing into dense asperate ones or granules towards sides.

Elytra about 1.39 to 1.47 times as wide as long (n=10), with eight striae including one
along epipleural margin; striae shallowly and finely impressed and ridged on both sides through¬
out; strial punctures sparse and very weak, without distinctly notching intervals; 7th stria almost
parallel to the 6th; intervals nearly flat, clearly micro-granulose, sparsely and finely punctate,
the punctures becoming coaser towards the outer intervals.

88

Teruo Ochi and Masahiro Kon

Pygidium carinate at base, weakly convex, micro-granulose, densely covered with trans¬
verse ocellate punctures. Prothorax with anterior angles shallowly hollowed on the ventral side.
Metasternum subtriangularly elevated in front, and then gently sloping downwards, with the
apex of the elevation produced anteriad and keeled along midline. Protibiae relatively slender,
curved inwards, armed with four strong teeth; terminal spur spatulate and rather elongate.

Aedeagus elongate. Phallobase about 2.2 to 2.5 mm in length (n=10), about 1.0 to 1.1 mm
in apical width (n=10). Parameres about 1.2 to 1.3 mm in length, each with two ventral teeth
from lateral aspect.

Female. Head with the median clypeal tooth more elongate and the basal incisions of the
tooth distinctly deeper; rugosities on clypeus stronger. Pronotum almost simple as in minor
males. Protibiae broader, with stronger lateral teeth.

Type series. Holotype: <?, Malaysia, Sabah State, Mt. Kinabalu, Sayap, 1200 m alt., 8. XI.

1994, T. Kikuta leg. Paratypes: 34 exs., Poring, Sabah State, Malaysia, 13. IV. 1995, T. Kikuta
leg.; 39 exs., Tahubang, Sabah State, 19. IV. 1995, T. Kikuta leg.; 6 exs., ditto, 20. IV. 1995;
65 exs., 24. II. 1995; 25 exs., ditto, 24. I. 1995; 61 exs., Sayap, Sabah State, Malaysia, 25. III.

1995, T. Kikuta leg.; 5 exs., ditto, 11. VIII. 1995; 12 exs., ditto, 7. XI. 1995.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. This species is named in honor of Mr. Francis Liew, Sabah Parks, who gave us
invaluable help for our researches in the Kinabalu National Park.

Notes. The present new species is closely related to Onthophagus sarawacus Harold
from Borneo, but can be distinguished from the latter by the following characters; 1) body more
strongly mat on the dorsal side; 2) head with clypeal margin strongly produced as a strong tooth
at the middle and clearly notched on both sides of the protrusion, whereas in O. sarawacus the
clypeal margin is also produced as a strong tooth but clearly not notched on either side of the
protrusion (cf. Fig. 9); 3) head with vertex strongly and widely carinate instead of being weakly
and rather briefly carinate; 4) in large indivisuals, pronotum with a distinctly depressed declivity
in front, whereas in O. sarawacus , it is almost simple or at most very slightly depressed; 5) in
the male genitalia, parameres rather long, each with two ventral teeth larger (cf. Fig. 12).

Onthophagus ( Parascatonomus) kikutai sp. nov.

(Figs. 3, 13-15)

Length: 4.6-8.5 mm; width: 3.5-4.4 mm (n=4).

Male. Body small-sized, oblong-oval, strongly convex above, and deeply constricted
between pronotum and elytra; dorsal side with head and pronotum shining and almost glabrous
except for the slightly setiferous sides of the latter, elytra almost opaque, somewhat sparsely
clothed with semi-recumbent short yellowish hairs; ventral side weakly shining, partly densely
clothed with yellowish hairs. Color brownish-black: head and pronotum suffused with blight
cupreous luster; elytra, pygidium, legs and ventral surface tinged with weak cupreous luster;
mouth organs, palpi, antennal foot-stalks reddish-brown; antennae yellowish-brown.

Head simple, distinctly transverse; clypeal margin rounded and weakly reflexed though
slightly truncated at the middle; clypeo-frontal suture completely effaced; genal suture fine, not

Six New Species and a New Subspecies of Onthophagus from Borneo

89

Figs. 13-16. Onthophagus ( Parascatonomus ) spp.-13, O. ( P .) kikutai sp. nov., male, head and

anterior part of pronotum, dorsal view; 14, right protibia, dorsal view; 15, aedeagus, dorsal and
lateral views.-16, O. ( P .) semicupreus Harold, aedeagus, dorsal and lateral views.

carinate; genae well produced laterally, rounded at apex; vertex also simple, without a carina or
tubercle; surface strongly, somewhat rugosely, and densely punctate on clypeus, the punctures
becoming smaller and sparser on the middle of clypeus and vertex. Antennae short and com¬
pact; scape short, invisible in dorsal view; club segments successively diminishing the size in
breadth and length.

Pronotum simple, strongly convex, about 1.24 to 1.33 times as wide as long (n=4); a medi¬
an longitudinal impression along midline almost imperceptible; anterior margin emarginated,
finely bordered throughout; lateral margins evenly rounded in front, clearly sinuate behind, fine¬
ly bordered; basal margin obtusely angulate at the middle and only slightly raised at the tip,

90

Teruo Ochi and Masahiro Kon

without distinct marginal line; anterior angles bluntly subangulate, rounded apically; posterior
angles obtuse; surface sparsely and finely punctate, the interspaces between punctures sparsely
punctulate and almost shining, the punctures becoming denser, coarser and a little asperate
towards sides.

Elytra about 1.24 to 1.35 times as wide as long (n=4), with eight striae including one along
epipleural margin; striae strongly and distinctly impressed, and ridged on both sides throughout;
strial punctures sparse and a little strong, with weakly notching intervals; 7th stria almost paral¬
lel to the 6th; intervals weakly convex, clearly micro-granulose, sparsely, finely, and somewhat
asperately punctate.

Pygidium well convex near apex, carinate at base, weakly shining though micro-granu¬
lose, densely covered with transverse ocellate punctures. Prothorax with anterior angles exca¬
vated on the ventral side. Metastemum subtriangularly elevated in front, and then sloping down¬
wards anteriad, with the apex of the elevation sharply defined. Protibiae moderately elongate,
curved inwards, armed with four strong teeth; terminal spur somewhat spatulate, not so short.

Aedeagus a little robust. Phallobase about 1.4 mm in length (n=l), about 0.6mm in apical
width (n=l). Parameres about 0.7 mm in length (n=l), each with a strong ventral tooth near
apex.

Female. Unknown.

Type series. Holotype: Poring, 800 m, Sabah State, Malaysia, 13. IV. 1995, T. Kikuta

leg. Paratypes: 1 c?., ditto, 17. V. 1995; 2 exs., Tahubang, Sabah State, 20. IV. 1995, T. Kikuta
leg.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. This species is named in honor of Mr. Toru Kikuta who made intensive
researches on dung beetles at Mt. Kinabalu.

Notes. The present new species is closely related to Onthophagus ( Parascatonomus) semi¬
cupreus Harold from the Sunda Islands, but can be distinguished from the latter by the follow¬
ing characters: 1) head simple, without two transverse carinae on frons and vertex, whereas in
O. semicupreus, it has two distinct carinae on frons and vertex; 2) elytra with intervals more
densely and more strongly punctate; 3) pygidium distinctly micro-granulose instead of being
shining and smooth to weakly shining; 4) male protibiae with rather elongate terminal tooth,
whereas in O. semicupreus , it is distinctly shorter and broader; 5) male genitalia is different in
shape (cf. Fig. 16).

Onthophagus (. Parascatonomus) anitidus sp. nov.

(Figs. 4, 17-18)

Length: 6.1-7.5 mm; width: 3.4-4.0 mm (n=238).

Body moderately convex, obviously constricted between pronotum and elytra; dorsal side
shining on head and pronotum, opaque on elytra, almost glabrous; ventral side shining, partly
clothed with pale white-yellowish hairs. Color usually uniformly black, with weak purplish lus¬
ter near anterior angles of pronotum; palpi, antennal foot-stalks reddish-brown, club segments
of antennae yellowish-brown.

Six New Species and a New Subspecies of Onthophagus from Borneo

91

Figs. 17-18: Onthophagus ( Parascatonomus ) anitidus sp. nov., male.-17, Head and anterior

part of pronotum, dorsal view; 18. aedeagus, dorsal and lateral views.

Male. Head distinctly transverse, subpolygonal in outline; clypeus subtrapezoidal, emar-
ginated at the middle, gently rounded on both sides, finely bordered; genae well produced later-
ad, obtusely angulate at the middle; front-clypeal suture weakly and finely carinate, a little
procurved; vertex with a barely perceptible weak carina in major male, often effaced in minor
males; surface weakly micro-granulose, densely and strongly punctate, the punctures becoming
smaller and sparser on vertex, transversely wrinkled on anterior part of clypeus. Antennae short
and compact; scape short, invisible in dorsal view; club segments successively diminishing the
size in breadth and length.

Pronotum simple, evenly convex dorsally, about 1.35-1.40 times as wide as long (n=5),
with a very slight longitudinal impression in basal fourth along midline; anterior margin emar-
ginated, finely bordered laterad, not distinctly so in the middle; lateral margins gently rounded
in front, sinuate behind, finely bordered; basal margin obtusely angulate at the middle and only
slightly raised at the tip, not distinctly bordered; anterior angles roundly subangulate, though

92

Teruo Ochi and Masahiro Kon

well produced forwards; posterior angles obtuse; surface usually shining, frequently opaque
weakly, sparsely and finely punctate, the interspaces between punctures sparsely punctulate, the
punctures becoming denser and coarser towards sides.

Elytra about 1.13 to 1.18 times as wide as long (n=4), with eight striae including one along
epipleural margin; striae shallowly impressed though rather wide and ridged on both sides
throughout; strial punctures sparse and very weak, with slightly notching intervals; 7th stria
weakly curved; intervals almost flat, clearly micro-granulose, sparsely and fairly finely punctate.

Pygidium convex near apex, carinate at base, weakly shining, densely covered with trans¬
verse ocellate punctures. Prothorax with anterior angles excavated on the ventral side. Meta¬
sternum subtriangularly elevated in front, and then sloping downwards anteriad, with the apex
of the elevation sharply defined. Protibiae moderately elongate, curved inwards, armed with
four strong teeth; terminal spur rather broad, briefly lanceolate, and decurved though well
toothed in dorsal view.

Aedeagus elongate. Phallobase about 1.3-1.4 mm in length (n=3), about 0.5 mm in apical
width (n=l). Parameres about 0.6 mm in length (n=l), each with a ventral tooth obtuse, not
strongly produced in lateral view though the tooth distinct in dorsal view.

Female . Head less transverse; clypeus a little strongly produced forwards; surface distinct¬
ly rugose on clypeus and frons; vertex more strongly and closely punctate. Elytra with intervals
more strongly micro-granulate.

Type series. Holotype: <?, Headquarter, Kinabalu Park, 1800 m, Sabah State, Malaysia,
12. II. 1995, T. Kikuta leg. Paratypes: 6 exs., the same data as the holotype; 2 exs., ditto, 13. X.
1994, T. Kikuta leg.; 3 exs., ditto, 21. X. 1994; 2 exs., ditto, X. 1994; 28 exs., ditto, 2. II. 1995;
6 exs., ditto, 12. II. 1995; 2 exs., ditto, 3. IV. 1995; 4 exs., ditto, 28. II. 1995; 11 exs., ditto, 2.
III. 1995; 12 exs., ditto, 3. III. 1995; 15 exs., ditto, 4. III. 1995; 2 exs., ditto, 1. V. 1995; 15 exs.,
ditto, 14. XI. 1997; 46 exs., ditto, 12. X. 1997; 20 exs., ditto, 27. VII. 1998; 12 exs., 23. XII.
1998; 8 exs., ditto, 24. XII. 1998; 3 exs., Liwagu, Sabah State, Malaysia, X. 1994, T. Kikuta
leg.; 1 ex., ditto, 4. IV. 1995; 2 exs., ditto, 6. IV. 1995; 15 exs., Sayap, nr. Kinabalu, Sabah
State, Malaysia, 12. V. 1995, T. Kikuta leg.: 13 exs., ditto, 25. III. 1995; 4 exs., ditto, 25. III.
1995; 5 exs., ditto, 11. V. 1995.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. The specific name means that the present new species is similar to Ontlio-
phagus ( Parascatonomus) nitidus Waterhouse.

Notes. The present new species is somewhat related to Onthophagus ( Parascatonomus )
semicupreus Harold from the Sunda Islands, but can be distinguished from the latter by the fol¬
lowing characters: 1) body uniformly black, whereas in O. semicupreus , head and pronotum
have strong cupreous luster; 2) head with one weak transverse carina, whereas in O. semi¬
cupreus it has two strong transverse carinae; 3) elytra with intervals flat instead of being weakly
convex; 4) parameres of male genitalia different in shape.

Six New Species and a New Subspecies of Onthophagus from Borneo

93

Onthophagus ( Parascatonomus) gunsalami sp. nov.

(Figs. 5, 19-22)

Length: 6.6-7.7 mm; width: 3.5^1.0 mm (n=3).

Body moderately convex above, rather shallowly constricted between pronotum and ely¬
tra; dorsal side shining, completely glabrous; ventral side shining, partly clothed with yellowish
hairs. Color uniformly black, frequently with slight puiplish luster on head and pronotum; palpi,
antennal foot-stalks reddish-brown, club segments of antenna yellowish-brown to dark yellow¬
ish-brown.

Male. Head somewhat transverse, subpentagonal in outline; clypeal margin subtrapezoidal
though slightly truncated at the middle, gently rounded on both sides, a little broadly bordered;
genae well produced laterad, obtusely angulate at the middle; front-clypeal suture clearly and
rather thickly carinate, the carina weakly arched forwards; vertex weakly and transversely
raised, though not distinctly carinate; surface shining, densely covered with small punctures on
vertex, rugosely punctate on frons and clypeus, densely and vaguely on ganae. Antennae short
and compact; scape short, invisible in dorsal view; club segments successively diminishing the
size in length and breadth.

Pronotum simple, evenly convex dorsally, rather wide, about 1.29-1.32 times as wide as
long (n=3), with a very slight longitudinal impression in basal half along median line; anterior
margin emarginated, finely bordered laterad, not distinctly so in the middle; lateral margins
evenly rounded in front, sinuate behind, finely bordered; anterior angles produced forwards,
rounded at tip; posterior angles obtuse; basal margin obtusely angulate at the middle, only
slightly raised at the tip, without a distinct marginal line; surface shining, moderately densely
and evenly punctate, the interspaces between punctures not distinctly punctulate, the punctures
becoming denser and coarser towards sides

Elytra about 1.21 to 1.32 times as wide as long (n=3); striae strongly and rather broadly
impressed and ridged on both sides throughout; strial punctures sparse and transverse, with
slightly notching intervals; 7th stria slightly curved; intervals weakly but clearly convex, shin¬
ing, sparsely and finely punctate.

Pygidium clearly convex in the middle, carinate at base, strongly shining, densely covered
with strong punctures though a little ocellate; apical margin fairly broadly bordered at the mid¬
dle. Prothorax with anterior angles excavated on the ventral side. Metasternum subtriangularly
elevated in front, and then sloping downwards anteriad, with the apex of the elevation obtuse,
not sharp. Protibiae a little elongate, incurved, armed with four strong teeth; terminal spur
broadened at the middle, briefly lanceolate, and decurved.

Aedeagus relatively small, elongate. Phallobase about 1.2 mm in length (n=l), about 0.5
mm in apical width (n=l). Parameres about 0.5 mm in length (n=l), each with a ventral tooth
sharp in lateral view.

Female. Head with clypeus more strongly produced forwards, and more densely rugose;
front-clypeal suture more thickly carinate than in male; vertex a little distinctly and transversely
raised. Protibiae with four external teeth stronger; terminal spur sharp and a little decurved.

Type series. Holotype: Poring 900 m, Sabah State, Malaysia, 16. I. 1997, T. Kjkuta

leg. Paratypes: 1 1 Poring, 1200 m, 13. IV. 1995, T. Kikuta leg.; 1 <?, ditto, 28. III. 1997,
T. Kikuta leg.

Type depository. The holotype is deposited in the collection of the Institute for Tropical

94

Teruo Ochi and Masahiro Kon

Figs. 19-22. Onthophagus (Parascatonomus) gunsalami sp. nov., male.- 19, Head and

pronotum, dorsal view; 20, right mesotibia, dorsal view; 21, right metatibia, dorsal view;
22, aedeagus, dorsal and lateral views.

Biology and Conservation, University Malaysia Sabah.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. This species is named in honor of Mr. Gnik Gunsalam who gave us invalu¬
able help for our researches in Sabah.

Notes. The present new species is closely related to Onthophagus ( Parascatonomus ) tami-
jii Kon, Ochi et Sakai from Borneo, but can be distinguished from the latter by the following
characters: 1) body much larger; 2) in the female, head with clypeal margin truncate at the mid¬
dle, whereas in O. tamijii , it is weakly bidentate; 3) pronotum less coarsely punctate; 4) elytra
with interval more coarsely punctate.

Six New Species and a New Subspecies of Onthophagus from Borneo

95

Onthophagus ( Parascatonomus ) sayapensis sp. nov.

(Figs. 6, 23-25)

Length: 4.1-6.1 mm; width: 2.0-3.1 mm (n=15).

Body moderately convex above, distinctly constricted between pronotum and elytra; dor¬
sal side shining, entirely glabrous; ventral side shining, partly clothed with yellowish hairs.
Color uniformly black to blackish-brown, sometimes partly with slight purplish luster; palpi,
antennal foot-stalks reddish-brown, club segments of antenna yellowish-brown.

Male. Head transverse, subpentagonal in outline; clypeus subtrapezoidal, slightly truncat¬
ed at the middle, gently rounded on both sides, finely bordered; genae well produced laterad,
obtusely angulate at the middle; front-clypeal suture clearly and a little finely carinate, the Cari¬
na slightly curved forwards, frequently forming an obtuse angle at the middle; vertex obtusely
and transversely raised, though not distinctly carinate; surface shining, densely and transversely
rugose or granulate on clypeus, closely and vaguely punctate or sculptured on the remaining
portions. Antennae short and compact; scape short, invisible in dorsal view; club segments suc¬
cessively diminishing the size in length and breadth.

Pronotum simple, evenly convex dorsally, about 1.24-1.37 times as wide as long (n=5),
with a very slight longitudinal impression in basal half along midline; anterior margin emar-
ginated, finely bordered laterad, not distinctly so in the middle; lateral margins well rounded in
front, sinuate behind, finely bordered; anterior angles produced forwards, rounded at tip; poste¬
rior angles obtuse; basal margin obtusely angulate at the middle, only slightly raised at the tip,
without distinct marginal lines; surface shining, sparsely and finely punctate, the interspaces
between punctures not distinctly punctulate as in the proceeding species, the punctures becom¬
ing denser and fairly coarser towards sides.

Elytra about 1.17 to 1.22 times as wide as long (n=4); striae rather strongly impressed and
ridged on both sides throughout; strial punctures sparse and distinct, with slightly notching
intervals; 7th stria almost parallel to the 6th; intervals weakly convex, usually strongly shining,
frequently very slightly micro-granulose, sparsely and finely punctate.

Pygidium clearly convex near apex, carinate at base, strongly shining, densely covered
with strong punctures though a little small; apical margin fairly broadly bordered. Prothorax
with anterior angles excavated on the ventral side. Metasternum subtriangularly elevated in
front, and then sloping downwards anteriad, with the apex of the elevation obtuse, not pointed.
Protibiae moderately elongate, curved inwards, armed with four strong teeth; terminal spur
broadened at the middle, briefly lanceolate, and decurved.

Aedeagus elongate. Phallobase about 1.0 mm in length (n=3), about 0.4 mm in apical
width (n=l). Parameres about 0.4 mm in length (n=l).

Female. Head less transverse; clypeus a little strongly produced forwards, surface distinct¬
ly rugose on clypeus and frons; vertex more strongly and closely punctate. Elytra with intervals
more strongly micro-granulate.

Type series. Holotype: d\ Sayap, nr. Kinabalu, Sabah State, Malaysia, 8. XI. 1995, T.
Kikuta leg. Paratypes: 11 exs., the same data as for the holotype; 1 <?, Headquarter, 1500 m,
Mt. Kinabalu, Sabah State, Malaysia, X. 1995, T. Kikuta leg.; 2 exs., Tahubang, nr. Kinabalu,
Sabah State, Malaysia, 24. II. 1995, T. Kikuta leg.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Biology and Conservation, University Malaysia Sabah.

96

Teruo Ochi and Masahiro Kon

Imm

Figs. 23-25. Onthophagus ( Parascatonomus ) sayapensis sp. nov., male.-23, Head and

anterior part of pronotum, dorsal view; 24, right protibia, dorsal view; 25, aedeagus,
dorsal and lateral views.

Distribution. Sabah State, Malaysia (Northern Borneo).

Etymology. The species is named after a place name, Sayap, near Mt. Kinabalu.

Notes. The present new species is closely related to Onthophagus (.Parascatonomus ) tami-
jii Kon, Ochi, et Sakai from Borneo, but can be distinguished from the latter by the following
characters: 1) elytra with striae more strongly and broadly impressed instead of being shallower
and finer; 2) elytra with intervals more strongly punctate, whereas in O. tamijii , they are very
finely punctate; 3) pygidium with apical margin more broadly bordered; 4) in the female, head
with clypeal margin truncated at the middle, whereas in O. tamijii , it is slightly but clearly emar-
ginated at the middle; 5) in the female, head with vertex clearly less transversely raised, whereas
in O. tamijii , it is distinctly and transversely raised; 6) in the female, head with clypeus more
strongly rugose on anterior portion. The present new species is also closely related to O. niasen-
sis Boucomont from the Nias Island, Indonesia, but can be easily distinguishable from the latter
by the finely bordered apex of pygidium, the elongate terminal spur of protibiae in the male, and
differently shaped male genitalia.

Six New Species and a New Subspecies of Onthophagus from Borneo

97

Onthophagus ( Parascatonomus ) katoi poringensis subsp. nov.

(Fig. 7)

Onthophagus ( Parascatonomus ) pilurarius : Balthasar, 1963, Monogr. Scarab., 2: 480; Kon et al., 2000,
Ent. Sci., 3: 371 (synonymy) [nec Lansberge 1883].

The present new subspecies differs from the nominotypical one from the Philippines as
follows: 1) body smaller (4.5-5.7 mm), whereas in the nominotypical species, the body is a little
larger (4.6-6.8 mm); 2) head with clypeus less produced forwards, and more strongly and close¬
ly rugose; 3) head with vertexal carina more distinctly carinate; 4) elytra with stria more widely
and strongly impressed, whereas in the nominotypical subspecies, it is rather finely and a little
shallowly impressed.

Length: 4.5-5.7 mm; width: 2.5-3.5 mm (n=15).

Type series. Holotype: d\ Poring, 900 m, Sabah State, Malaysia, 9. I. 1998, T. Kikuta
leg. Paratypes: 13 exs., the same data as for the holotype; 2 exs., Mt. Bawang, Kalimantan,
Indonesia, IX. 1990; 1 ex., ditto, VIII. 1990.

Type depository. The holotype is deposited in the collection of the Institute for Tropical
Botany and Conservation, University Malaysia Sabah.

Distribution. Borneo.

Etymology. The species is named after a place name, Poring, Sabah State.

Notes. The distributional records of Onthophagus ( Parascatonomus ) pilurarius Lans¬
berge from Borneo may be those of O. ( P .) katoi poringensis subsp. nov. We have not examine
the specimens of the true O. (P.) pilurarius from Borneo.

Acknowledgments

We wish to express our cordial thanks to Messrs. M. Fujioka and T. Kikuta for giving us
the opportunities of examining invaluable specimens. This study was supported in part by a
Grant-in-Aid from the Japan Society for the Promotion of Science (No. 14405013).

m fa

• i£ #1$ : JlCTt(il§8$|x) — tf )\

—LT,

H, ParascatonomusMM^ 6 frtt, Onthophagus (Parascatonomus) riekoae sp. nov., O. (P.)
liewi sp. nov., O. (P.) kikutai sp. nov., O. (P.) anitidus sp. nov., O. ( P .) sayapensis sp. nov.,
O. (P.) gunsalami sp. nov. Jo ± If O. ( P .) katoi Ochi et Kon <D 1 O. (P.) katoi poring¬

ensis subsp. nov. IS® L tz .

References

Balthasar, V., 1963. Monographic der Scarabaeidae und Aphodiidae der palaearktischen und orientalis-
che Region, 2: 1-628. Prag.

Boucomont, A., 1914. Les Coprophages de l'Archipel Malais. Annales de la Societe Entomologique de
France , 83: 238-350.

98

Teruo Ochi and Masahiro Kon

Boucomont, A. and J. Gillet, 1921. Faune entomologique de l'lndochine fran 9 aise. Fam. Scarabaeidae
(Laparosticti): 1-76. Saigon.

Harold, E., von, 1877. Enumeration des Lamellicomes Coprophages rapportes de l'Alchipel Malais, de la
Nouvelle Guinee de l'Austrailie boreale par M. M. J. Doria, O. Beccari et L. M. D' Albertis, par le
Baron E. de Harold. Annali Museo civico di Storici naturale di Genova , 10: 38-109.

Kon, M., S. Sakai and T. Ochi, 2000. A new species of the genus Onthophagus (Coleoptera: Scara¬
baeidae) from Sarawak, Borneo. Entomological Science, 3: 367-371.

Nomura, S., 1976. On the subgenus Parascatonomus from Japan and Taiwan. The Entomological Review
of Japan, 29: 25-33.

Ochi, T. and K. Araya, 1992. Studies on the coprophagous scarab beetles from East Asia. II (Coleoptera,
Scarabaeidae). Giornale italiano di Entomologia, 6: 79-108.

-and-, 1996. Studies on the coprophagous scarab beetles from East Asia. Ill (Coleoptera,

Scarabaeidae). Giornale italiano di Entomologia, 8: 1-15.

Paulian, R., 1945. Coleopteres Scarabeides de l'lndochine. Faune de TEmpire Francaise, 3: 1-225.
Larose, Paris.

Sharp, D., 1875. Descriptions of some new genera and species of Scarabaeidae from tropical Asia and
Malaysia, Part II. Coleopterologische Hefte, 14: 47-66.

(Received March 18, 2005; Accepted April 11, 2005)

99

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The Entomological Review of Japan Vol. 60, No. 1

CONTENTS

Hayashi, M. and O. Tominaga: Records of Donaciinae from Primorsky Province in 2004, with Notes on

the Distribution of Plateumaris shircihcitcii Kimoto (Coleoptera: Chrysomelidae).. 1

Morita, S.: A New Pterostichus (Coleoptera: Carabidae) from the Islands of Tsushima, West Japan.. 9

Hoshina, H. and M. Sato: Synonymic Notes on Two Species of the Families Hydrophilidae and Leiod-

idae (Coleoptera) from Japan..13

Ashida, H.: The Complex of Trechiama fujitai (Coleoptera: Trechinae) from Hyogo Prefecture, West
Japan (III) — A New Relative of Trechiama latilobatus Ashida —.. 17

Tominaga, O., Y. Imura, M. Okamoto, Z.-H. Su, T. Ojika, N. Kashiwai and S. Osawa: Origin of
Ohomopterus uenoi (Coleoptera: Carabidae: Carabinae) as Deduced from Comparisons of DNA
Sequences of Mitochondrial ND5 Gene and Nuclear Internal Transcribed Spacer I (ITS I) with

Morphological Characters..23

Imura, Y., K. Akita, M. Okamoto, O. Tominaga, N. Kashiwai, Z.-H. Su, T. Ojika and S. Osawa: On
Ohomopterus arrowicinus kirimurai (Coleoptera: Carabidae) as Examined by Phylogenetic Trees of

Mitochondrial ND5 Gene and Nuclear ITS I DNA as well as Morphology of Genital Organs..35

Ito, N.: Five New Species of the leptopus Group of Harpaline Genus Trichotichnus (Coleoptera: Carab¬
idae) from Central and Northeastern Japan, with Note on Taxonomic Position of T. tsurugiyamanus.

.39

Ito, N.: Replacement Names for the Junior Homonyms of Two Species of Trichotichnus and a Genus of

Harpalini (Coleoptera: Carabidae: Harpalini)..53

Arimoto, H. and S. Riese: Propsephus nanshanus , a New Species of Elateridae (Coleoptera) from Taiwan.

.55

Narukawa, N. and H. Ashida: A New Subspecies of Episcaphula matsumurai Chujo (Coleoptera:

Erotylidae) from the Yaeyama Group in the Southern Ryukyus, Southwestern Japan..59

Hayashi, Y.: A New Genus and Species of Coprophilini (Coleoptera: Staphylinidae: Oxytelinae) from

Japan..63

Hayashi, Y.: New Records of Staphylinidae from Taiwan, 4..68

Ochi, T. and M. Kon: Notes on the Coprophagous Scarab-beetles (Coleoptera: Scarabaeidae) from

Southeast Asia (VI) —Three New Taxa of the Tribe Coprini from Borneo —..69

Ochi, T. and M. Kon: Notes on the Coprophagous Scarab-beetles (Coleoptera: Scarabaeidae) from

Southeast Asia (VII) — Three New Species of Onthophagus (Phanaeomorphus) from Borneo —..75

Ochi, T. and M. Kon: Notes on the Coprophagous Scarab-beetles (Coleoptera: Scarabaeidae) from
Southeast Asia (VIII) — Six New Species of Onthophagus ( Parascatonomus ) and a New Subspecies of O. (P.)
katoi from Borneo —..83

99